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Journal of

Hymenoptera
Research

Volume 15, Number 1 April 2006

ISSN #1070-9428
CONTENTS

GIBSON, G. A. P. A revised concept of Spalangia philippinensis Fullaway, 1917 (Hymenoptera:

Pteromalidae) 1

KIMSEY, L. S. and M. S. WASBAUER. Phylogeny and checklist of the nocturnal tiphiids of the

Western Hemisphere (Hymenoptera: Tiphiidae: Brachycistidinae) 9

KUHLMANN, M. Fauna and biogeography of the bees and wasps of the Cook Islands

(Hymenoptera Aculeata) 26

PINTO, J. D. A review of the New World genera of Trichogrammatidae (Hymenoptera) 38

PULAWSKI, W. J. Nomenclatural changes in Old World Crabronidae (Hymenoptera), with

taxonomic comments and new distribution records 164

PUNZO, F. Plants whose flowers are utilized by adults of Pepisis grossa Fabricius (Hymenoptera:

Pompilidae) as a source of nectar 171

NOTE:

STARR, C. K. and A. W. HOOK. Polistes goeldii (Hymenoptera: Vespidae) is a widespread

but rare social wasp 177

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J. HYM. RES.
Vol. 15(1), 2006, pp. 1-8

A Revised Concept of Spalangia philippinensis Fullaway, 1917

(Hymenoptera: Pteromalidae)

Gary A. P. Gibson

Agriculture and Agri-Food Canada, Biodiversity and Integrated Pest Management, K. W. Neatby

Bldg., 960 Carling Avenue, Ottawa, Ontario, Canada, K1A 0C6

"Address for correspondence: Dr. Gary Gibson, K.W. Neatby Building, 960 Carling Avenue,

Ottawa, Ontario, Canada K1A 0C6; e-mail: gibsong@agr.gc.ca; tel 613-759-1823; fax 613-759-1927

Abstract. — A lectotype male and three paralectotype females are designated for Spalangia
philippinensis Fullaway, 1917, which is removed from synonymy with S. endius Walker, 1839, and
placed in synonymy with S. cameroni Perkins, 1910, syn. nov. Two of the female paralectotypes are
conspecific with S. endius, whereas the third is conspecific with S. gemina Boucek, 1963. The revised
concept and new synonymy are based on comparison of the original description and illustrations of
S. philippinensis with the four specimens that are interpreted as syntypes. The lectotype is selected
so as to least disrupt current nomenclature while being compatible with the original description.
Current concepts of S. cameroni and S. simplex Perkins, 1910, as interpreted by Boucek (1963), are
confirmed by examination of type material of these species.

Fullaway (1917) described and illustrat-
ed both sexes of Spalangia philippinensis
(Hymenoptera: Pteromalidae) from a cul-
ture that had been established in 1914 from
house fly, Musca domestica L. (Diptera:
Muscidae), puparia and other muscid
puparia collected in the Philippines. The
parasitoids were propagated and released
in Hawaii as part of a control program for
the horn fly, Haematobia irritans (L.) (Dip-
tera: Muscidae). Fullaway briefly com-
pared his new species to S. cameroni
Perkins, 1910, but did not state the number
of females and males he had before him,
nor select a holotype or state where type
material was deposited.

Boucek (1963) revised the world species
of Spalangia Latreille. In this work he
synonymized S. philippinensis under S.
endius Walker, 1839, based on two females
that Fullaway had sent him "of his
species", and newly described S. geniina
based on females and males from Maur-
itius, India, Thailand, Malaysia, Fiji, and
Venezuela. The name S. philippinensis has
not been used in any scientific publication

since Boucek (1963) except as a synonym of
S. endius or in simple lists of taxa. In
contrast, extensive information has been
published under the name S. geniina, in-
cluding research on its life history (Morgan
et al. 1989, 1991), biological attributes
(Costa 1995; Geden 1999, 2002) and host-
parasitoid modelling (Geden 1996, 1997).
Several publications also compare it to
other Spalangia species or list new distri-
bution and host records (see Noyes 2003
for summary), and three partial sequences
of the 12S and 28S ribosomal RNA genes
have been deposited in GenBank under the
name S. geniina (accession numbers
AF289673, AY855200 and AY8500201).

As part of research investigating the
identity and diversity of chalcid parasi-
toids of filth-breeding flies in North Amer-
ica, I borrowed type material of three
species of Spalangia housed at the Bernice
P. Bishop Museum, Honolulu, Hawaii
(BPBM). The material included a single
female syntype of S. cameroni (labelled as
holotype no. 1578), a male and female
syntype of S. simplex Perkins, 1910

Journal of Hymenoptera Research

(mounted together on one card and la-
belled as holotype no. 1579), and a female
labelled "type" and a male labelled "type
6A" of S. philippinensis. The latter two
specimens were mounted separately on
square cards1 and additionally bore three
identical printed labels with "Honolulu
H.T.", "D.T. Fullaway collector" and "In-
sectary".

Boucek examined the male type of S.
endius in The Natural History Museum,
London (BMNH), but he did not have the
opportunity to examine type material of S.
cameroni, S. philippinensis or S. simplex. My
study of the type specimens of S. cameroni
and S. simplex confirmed Boucek's (1963)
interpretation of these two names. Howev-
er, examination of the BPBM specimens
labelled as type and male type of S.
philippinensis revealed that the female is
conspecific with S. gemina and the male is
conspecific with S. cameroni. Following this
discovery, I investigated whether collec-
tions of the Department of Entomology,
University of Hawaii, Manoa (UHM) and
the Hawaii Department of Agriculture,
Division of Plant Industry, Honolulu
(HDOA) possessed any other potential
syntypes of S. philippinensis. The UHM
collection contains no specimens identified
as S. philippinensis, but I received three
females labelled as S. philippinensis from
HDOA. One of the HDOA females bears
three labels with "Honolulu Oahu", "orig-
inal cotype" and "Type material Spalangia
philippinensis". The latter label is of the
same red paper and Spalangia philippinensis
is in the same handwriting as the type
labels of the two BPBM specimens. The
female is mounted upside down on a plas-
tic point so that its propodeum is con-
cealed, and it lacks its head and antennae,
but sculpture of the pronotum indicates it
is a S. endius female. The other two HDOA

The female was partly broken and detached from the card
except by the apex of one fore wing when it arrived; I therefore
point-mounted it and pinned below it the original card with
pieces of the specimen still attached.

females are also S. endius. One is point-
mounted and bears three printed labels
with almost the same data as the BPBM
specimens, "Honolulu Oahu", "D.T. Full-
away collector", and "Insectary". The
other female bears a single handwritten
label with "Honolulu Oahu"; it is also
point-mounted but has a minuten pin
through the point that is pinned into
a circular piece of paper pierced by
a second, larger pin. Consequently, three
specimens are labelled variously as
"types" of S. philippinensis and these
comprise three different species — S. endius
Walker, S. cameroni Perkins, and S. gemina
Boucek. I therefore studied the original
description and illustrations in Fullaway
(1917) (dorsal habitus of female S. philippi-
nensis and female and male antenna of S.
philippinensis and S. cameroni) in an attempt
to determine which species, under current
concepts, was described as S. philippinensis.
The purpose of this paper is to select
a lectotype for S. philippinensis that least
affects stability of existing nomenclature
while still being compatible with the
original concept and description of Full-
away (1917).

MATERIALS AND METHODS

My concepts of S. gemina and S. endius
are based on Boucek (1963). Terms for
structure follow Gibson (1997). Relative
measurements were taken with a Nikon
SMZ-1500 microscope fitted with a 10 mm
ocular grid having 100 divisions. Scanning
electron photomicrographs of type-series
specimens of S. philippinensis were taken
using an environmental SEM and digitally
retouched using Adobe Photoshop™ in
order to enhance clarity.

RESULTS

Gibson (2000) provided an illustrated
key that differentiates S. cameroni, S. gemina
and S. endius from other introduced and
native species of Spalangia that are para-
sitoids of filth flies (Diptera: Muscidae) in
North America. Features used to differen-

Volume 15, Number 1, 2006

0imta «m«HNW»

Figs 1-3. 1, Spalangia philippinettsis, original habitus drawing adapted from Fullaway (1917). 2, S. endius, dorsal
mesosoma. 3, S. philippinensis, BPBM female paralectotype, dorsal mesosoma. (Abbreviations: fre = frenum,
no3 = metanotum, pits = anterior cells of paramedian crenulate furrows.) Scale bar = 200 u.

Journal of Hymenoptera Research

tiate S. endius from S. cameroni and S.
gemina include differences in sculpture of
the pronotum and propodeum, and a dif-
ference in the setal pattern of the fore wing
of males. Spalangia endius has the pronotum
comparatively sparsely punctate with cir-
cular depressions separated by flat, shiny
interspaces (Fig. 2), whereas S. cameroni
and S. gemina belong to a group of species
that share a rugulose-reticulate pronotum,
the depressions being so crowded as to be
irregular (Fig. 3). There is also a conspicu-
ous difference in propodeal sculpture
between S. endius and the other two
species. Although all three species have
a transverse row of crenulae along the
anterior margin of the propodeum (Figs 2,
3), in S. endius the paramedian crenulate
furrows posterior to the transverse row of
crenulations are parallel or only slightly
and evenly widened anteriorly so as to
form a narrowly V-shaped sculptural com-
plex. Furthermore, the anterior most cells
of the paramedian crenulate furrows are
similar in size to the other cells of the
furrows (Fig. 2). In contrast, both S.
cameroni and S. gemina have the parame-
dian crenulate furrows obviously widened
anteriorly so as to form more of a Y-like
sculptural complex, with the anterior most
cell or cells on either side of the median
carina being conspicuously larger than the
more posterior cells and usually also
tapered posteriorly (Figs 3, 4). Based on
these two features, both the original de-
scription and the female habitus illustra-
tion of S. philippinensis given by Fullaway
(1917) (reproduced here as Fig. 1) demon-
strate that the species could be S. gemina
and /or S. cameroni, but not S. endius.
Although the original female habitus illus-
tration is not detailed, the pronotum is
obviously coarsely sculptured (Fig. 1). Fur-
thermore, the description of the female
states "pronotum rugose and hairy",
which accurately describes the pronotum
of the BPBM female (Fig. 3) and male, but
does not correctly describe the pronotum
of any of the three HDOA females (Fig. 2).

Figs 4-8. 4, Spalangia philippinensis, male lectotype
propodeum. 5-8 antenna. 5, S. philippinensis, BPBM
female paralectotype (= S. gemina). 6, S. philippinensis,
HDOA female paralectotype (= S. endius). 7, S.
philippinensis, male lectotype (= S. cameroni). 8, S.
gemina, male. Scale bar = 200 u.

Volume 15, Number 1, 2006

Figs 9-11. Line drawings of antennae reproduced
from Fullaway (1917). 9, Spalangia philippinensis
female. 10, S. philippinensis male. 11, S. cameroni male.

The female description also states "a
longitudinal carina divides it [propo-
deum], in front passing between two rather
large shallow pits and behind flanked on
either side by a punctate line or furrow".
The female habitus clearly illustrates the
described pits as a triangular region (Fig. 1,
pits) posterior to the scutellar frenum and
metanotum (Fig. 1, fre, no3), similar to that
of the BPBM female (Fig. 3) and male
(Fig. 4) but not to the propodeal sculpture
of the HDOA females (Fig. 2). Finally,
although the fore wing is entirely bare
within the basal one-third of both sexes of
S. gemina and S. cameroni, males of S. endius
have conspicuous setae within and de-

fining the basal cell (see figures in Gibson
2000). For S. philippinensis the female fore
wing was described as "ciliate outwardly
from the juncture of submarginal with
marginal but basally bare" (Fig. 1). The
male of S. philippinensis was described only
relative to how it differed from the female
and the description does not mention any
difference in fore wing setation between
the sexes; therefore, a basally bare fore
wing must also be assumed for the male.
This represents a third feature that indi-
cates Fullaway was describing either S.
cameroni or S. gemina but not S. endius as S.
philippinensis.

In his key to species, Boucek (1963)
differentiated S. gemina from S. cameroni
based on slight differences in head shape
and antennal structure. Females of S.
cameroni were stated to have the second
funicular segment oblong and the distal
segments quadrate, whereas females of S.
gemina were stated to have the second
funicular segment subquadrate and the
following segments transverse. The female
description of S. philippinensis states "1st
funicular joint about equal to pedicel, the
next two joints about as broad as long, the
four following ones a trifle wider than
long". This accurately describes the anten-
na of the BPBM female labelled as type
(Fig. 5) and the female antenna Fullaway
illustrated as S. philippinensis (reproduced
here as Fig. 9), but conflicts with the
antennal structure of female S. endius.
Females of S. endius have the first funicular
segment obviously shorter than the pedicel
(Fig. 6), a fourth feature that suggests
Fullaway's concept of S. philippinensis was
not in the sense of S. endius. Boucek (1963)
further stated that males of S. cameroni
have the distal funicular segments clearly
oblong compared to hardly longer than
broad for males of S. gemina. The male
description of S. philippinensis states "the
first funicle joint long ...and the other
funicular segments all longer than wide".
The BPBM male syntype has all the
funicular segments obviously oblong

6 Journal of Hymenoptera Research

(Fig. 7), which is more similar to the male domestica as well as other muscid puparia
antenna that Fullaway illustrated as S. collected in the Philippines. It is therefore
cameroni (reproduced here as Fig. 11) than quite likely that his colony, and the type
to the original illustration provided for the series of S. philippinensis taken from it, was
male of S. philippinensis (reproduced here composed of more than one species. The
as Fig. 10) or to the antenna characteristic BPBM female labelled as "type" and the
of males of S. gemina (Fig. 8). Boucek (1963) male labelled as "type S" of S. philippinensis,
also described a difference in genal length which are also labelled "Insectary", show
between females of S. cameroni and S. that the colony consisted of at least two
gemina. In S. cameroni females the gena species, S. gemina and S. cameroni, respec-
was described as being slightly longer than tively. Under material examined for S.
the relatively small eyes, whereas in S. endiits, Boucek (1963) listed the data "Ha-
gemind it was said to be shorter than the waii: Philippine Spalangia" for the two
width of the eye. This difference was females that Fullaway sent to him as his
restated by Gibson (2000), who also de- species. Consequently, the specimens may
scribed males of S. cameroni as having the have been adults collected in the Philip-
gena only slightly less than the width and pines rather than cultured specimens and if
at least two-thirds the length of an eye, in so are not part of the original type series and
contrast to males of S. gemina, which have therefore ineligible for lectotype designa-
the gena distinctly less than the width and tion. However, the HDOA female labelled
less than half the length of an eye. The as "original cotype" and with a red type
original description of the male of S. label similar to the BPBM specimens is S.
philippinensis states that the head is shorter endius, and because of its labels I accept it as
than for the female, but no mention is a third syntype of S. philippinensis. I also
made of the length of the gena, which for interpret as a syntype the HDOA female
the female is described as "cheeks flat and with almost the same labels and data as the
as long as the eyes". This latter statement BPBM specimens. I exclude from the type
suggests that a female of S. cameroni rather series the HDOA female that has only
than S. gemina was being described as S. a single handwritten label because there is
philippinensis, but it conflicts with the no indication that it was a Fullaway spec-
description and illustration of the female imen or that it originated from the insectary.
antenna and is the only statement in the Regardless, the four remaining specimens I
female description that does not accurately interpret as syntypes indicate that the
reflect the BPBM female labelled as type, colony from which S. philippinensis was
In this female, relative measurements of described consisted of at least S. cameroni,
eye width: eye length: maximum genal s. gemina and S. endius. Therefore, designa-
length = 47 : 62 : 50. The same relative tion of a lectotype is necessary to fix the
measurements for the BPBM male are meaning of S. philippinensis.
39 : 51 : 40. jfoe current International Code of Zoo-
logical Nomenclature (ICZN 1999) recom-

DTSCl JSSTOM

mends that to preserve stability of nomen-

Multiple species of Spalangia are com- clature an author should act consistently

monly reared as part of surveys of pupal with or at least give great weight to

parasitoids of filth-breeding flies. Sulaiman previously accepted taxonomic restrictions

et al. (1990) reared S. cameroni, S. endius and of the application of the name when

S. gemina along with another species, S. designating a lectotype (recommendation

uigroaenea Curtis, 1839, in a survey in 74A). For S. philippinensis, such stability

peninsular Malaysia. Fullaway established could be achieved by designating one of

his colony from the puparia of Musca the two HDOA syntype females as lecto-

Volume 15, Number 1, 2006

type, which would retain Boucek's (1963)
synonymy of S. philippinensis under S.
endius. However, fixation of the name in
the sense of S. endius would be demonstra-
bly incorrect based on the original de-
scription and illustrations of S. philippinen-
sis provided by Fullaway (1917). I consider
such an obviously incorrect nomenclatural
action as inappropriate, even though Full-
away also misinterpreted his species many
years after the original description when he
sent specimens to Boucek. The original
description and illustrations clearly dem-
onstrate that one or both of S. cameroni and
S. gemina were described as S. philippinen-
sis. Fullaway's brief comparison of his new
species with S. cameroni demonstrates that
he was aware of the latter species and
considered that it was distinct, but it is
unclear whether the differential features he
gave (stouter antenna with first funicle
joint more or less obconic, and shorter
club) referred to the female of S. philippi-
nensis only. There are also discrepancies in
the descriptions and illustrations that
suggest these might have been from
a mixed series of S. cameroni and S. gemina.
The description of the female antenna
matches that of S. gemina and the BPBM
female labelled as type, whereas the de-
scription of the cheeks suggests a female of
S. cameroni was being described. The
illustration of the male antenna of S.
philippinensis (Fig. 10) is certainly more
characteristic of S. gemina (Fig. 8) than S.
cameroni (Fig. 7), but the corresponding
description states that the funicular seg-
ments beyond the first segment are all
"longer than wide", which is characteristic
of S. cameroni and certainly descriptive of
the antenna of the BPBM specimen labelled
as male type of the species (Fig. 7). There is
no information in the original publication
concerning whether the illustrations were
prepared from the actual specimens used
by Fullaway to prepare the descriptions, or
from other colony specimens, and it is
certain that the colony consisted of a mixed
culture. The original description and illus-

trations clearly demonstrate that S. philip-
pinensis was established in the sense of S.
gemina or S. cameroni, but it can not be
stated unequivocally that it was in the
current sense of only one of these two
names. Although the description of the
female is the primary description for the
species and the BPBM female is labelled as
"type", no holotype was selected in the
original description. I therefore designate
the BPBM male, labelled as "type J,
Spalangia philippinensis" , as lectotype of S.
philippinensis. I designate the correspond-
ing BPBM female and the two HDOA
females discussed above as paralectotypes
and have added my lectotype and para-
lectotype labels to the respective speci-
mens. The selection of the BPBM male as
lectotype is at least compatible with the
original description of S. philippinensis. It
also minimizes disruption of current no-
menclature because it retains the name as
a junior synonym and avoids the synony-
my of S. gemina Boucek, a name with an
extensive modern literature, which would
result if the BPBM female was selected as
lectotype of the species. I therefore remove
S. philippinensis Fullaway, 1917, from syn-
onymy with S. endius Walker, 1839, and
newly synonymize the name with S.
cameroni Perkins, 1910, syn. nov.

ACKNOWLEDGEMENTS

I thank Dr Gordon Nishida for the extended loan of
the BPBM Spalangia type material, Dr Bernarr Kuma-
shiro for the HDOA specimens, and Dr Dick Tsuda for
information concerning the UHM collection. I also
thank Ms Jennifer Read (AAFC, Ottawa) for the
scanning electron mircrographs and reproduction of
Fullaway's illustrations, and Dr John Huber (Canadi-
an Forest Service, Ottawa) and Dr James O'Hara
(AAFC, Ottawa) for reviewing and providing con-
structive criticism of an earlier version of this
manuscript.

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England.

Sulaiman, S., B. Omar, S. Omar, J. Jeffery, I. Ghauth,
and V. Busparani. 1990. Survey of microhyme-
noptera (Hymenoptera: Chalcidoidea) parasitiz-
ing filth flies (Diptera: Muscidae, Calliphoridae)
breeding in refuse and poultry farms in peninsu-
lar Malaysia. Journal of Medical Entomology 27:
851-855.

Walker, F. 1839. Monographia Chalciditum. 2. London.

J. HYM. RES.

Vol. 15(1), 2006, pp. 9-25

Phylogeny and Checklist of the Nocturnal Tiphiids of the Western
Hemisphere (Hymenoptera: Tiphiidae: Brachycistidinae)

Lynn S. Kimsey and Marius S. Wasbauer

Bohart Museum of Entomology, Department of Entomology, University of California, One Shields
Ave., Davis, CA, USA 95616, email: lskimsey@ucdavis.edu

Abstract. — Phylogenetic relationships among eleven genera of Brachycistidinae (Tiphiidae) .are
discussed, based on male characteristics, and a key to the genera is provided. The genus Stilbopogon
is found to be the sister group of the rest of the brachycistidine genera. There are two well
supported clades: Brachymaya + (Hadrocistis + Brachycistina) and (Paraquemaya + (Bradiycistellus +
(Dolichetropis + (Sedomaya + (Brachycistis + (Acanthetropis + Colocistis)))))). A complete synonymic
checklist of genera and species is also given, with new synonymy of the Cameron species
Brachycistis tysias, Brachycistis dubitatus and Brachycistis sciron under Brachycistis alcanor
(Blake), and Brachycistis fulvilabris and Brachycistis arines under Brachycistis vigilax Cameron.
In addition, Brachycistis ioachinensis Bradley is also newly synonymized under vigilax.

Key words. — Tiphiidae, Brachycistidinae, Acanthetropis, Brachymaya, Brachycistis, Brachycistellus,
Colocistis, Dolichetropis, Hadrocistis, Paraquemaya, Sedomaya, Stilbopogon

Nocturnal tiphiids in the Western Hemi- males have distinctive and informative

sphere all belong to the subfamily Brachy- modifications of the head, thorax, wing

cistidinae, which only occurs in the Amer- venation and genitalia. Most of the taxon-

icas. In the Palearctic and Afrotropical omy of Brachycistidinae is based on males.

Regions nocturnal tiphiids are all members Some aspects of the wing venation used to

of the subfamily Myzininae. There has discriminate species and genera, such as

never been a monographic review of the the number of submarginal or discoidal

Brachycistidinae, although a number of the cells, seem correlated with body size rather

genera have been at least partly revised than higher taxon. The greatest taxonomic

(Kimsey and Wasbauer 1998a, 1998b, Was- problem in this group is one shared by the

bauer 1958, 1966, 1968a). Brothers (1975), Mutillidae: males and females are adapted

Brothers and Carpenter (1993) and Kimsey for entirely different microhabitats and

(1991) gave Brachycistidinae status as the share few if any characteristics. Females

sister group of Tiphiinae. This monophy- are wingless and ant-like, lacking even

letic group is characterized by a number of ocelli. There are very few female brachy-

distinctive modifications, including in the cistidines in collections, as they are noctur-

male the pronotum lacking a dorsal sur- nal and are not collected by common

face, edentate tarsal claws, ocelli generally trapping methods. In Brachycistidinae this

enlarged, and forewing with abbreviated dichotomy has resulted in entirely differ-

marginal and submedial cells, and in the ent species and generic level nomenclature

female a single midtibial spur. for males versus females. Mickel and

Brachycistidine wasps are thought to be Krombein (1942) described seven genera

difficult taxonomically because they lack on the basis of females and there are

distinctive coloration and the species ap- currently 11 genera based on males. Was-

pear structurally similar. However, the bauer (1968b, 1971) associated some of the

Journal of Hymenoptera Research

Table 1. Character matrix for the brachycistidine genera based on male characteristics discussed in the text.

1 1

9 0

1 2

Tiphia

0 0

Stilbopogon

1 0

0 0

Brachymaya

0 0

Hadrocistis

0 0

Brachycistina

0 0

Acanthetropis

1 1

Colocistis

1 0

Brachycistis

1 0

Brachycistellus

1 0

Dolichetropis

1 0

Paraquemaya

0 0

Sedomaya

1 0

type species of these female-based genera
with males in the genus Brachycistis.
Additional synonymies were made by
Kimsey (2006). Clearly, more sex associa-
tions may be needed to eliminate extrane-
ous generic and species names in this
group, but this can probably only be
accomplished by more intensive collecting
efforts, DNA homologies, or by rearing
once hosts are known. A complete syno-
nymic checklist of the Brachycistidinae is
given below.

MATERIALS AND METHODS

Specimens used in this study came from
the Bohart Museum of Entomology, Uni-
versity of California, Davis (UCDC); Ento-
mology Research Museum, University of
California, Riverside (UCRC); the Califor-
nia Academy of Sciences, San Francisco
(CAS), and the personal collection of M. S.
Wasbauer. The species examined for this
study are indicated in the checklist by an
asterisk (*). Collection repositories cited in
the checklist include: MCZ - Museum of
Comparative Zoology, Harvard Universi-
ty, Cambridge, Massachusetts (S. Shaw);
CU - Department of Entomology, Cornell
University, Ithaca, New York (J. Liebherr);
SMEK Snow Entomological Museum,
University of Kansas, Lawrence (R.
Brooks); EMUS -■ Entomology Collection,
Utah State University, Logan (T. Gris-

wold); BMNH - The Natural History
Museum, London, S. Lewis; LACM - Los
Angeles County Museum of Natural His-
tory, Los Angeles, California (R. R. Snel-
ling); AMNH - American Museum of
Natural History, New York. City, New
York (J. M. Carpenter); ANSP - Philadel-
phia Academy of Sciences (D. Azuma);
CAS - California Academy of Sciences, San
Francisco (N. Penny); UMSP - University of
Minnesota, St. Paul (J. C. Luhman); and
NMNH - U. S. National Museum of
Natural History, Smithsonian Institution,
Washington, D. C. (A. S. Menke, K. V.
Krombein). The Cameron Brachycistis types
were borrowed from the Natural History
Museum, London.

Male characters below were prepared
using Winclada version 1.00.08 (Nixon
2002) and analyzed in Hennig86 (Farris
1988) and NONA version 2.0 (Goloboff
1999). The data matrix consisted of twenty-
three male characters and 13 taxa (Table 1).
The genus Tiphia was used to represent the
sister group Tiphiinae as the outgroup.
Species used in the analysis were selected
to give the widest range of characteristics
possible. However, once characters were
coded it was found that multiple species
from each genus were redundant and were
therefore eliminated from the analysis as
uninformative, resulting in a single exem-
plar species for each genus.

Volume 15, Number 1, 2006

Clinracters. —

1. Antennal socket: rim simple (0), with
ventral carinule or dilation (1). In the
majority of tiphiids the antennal sock-
ets are rimmed by a simple carina.
This is also the case in Stilbopogon
(Fig. 11). Therefore a simple antennal
socket carina is considered primitive.
In the other brachycistidine genera the
socket carina has a ventrally pointing
"tail", dilation or subsidiary carinule
(Fig. 7).

2. Eye margin: flattened or evenly con-
vex (0), emarginated (1). In Tiphia and
the majority of brachycistidine genera
the inner margin of male eye is
flattened or convex. The remaining
brachycistidine genera have the inner
eye margin indented or emarginate (as
in Fig. 7).

3. Clypeal shape: flat in profile (0),
convex (1) or concave (2). In tiphiines
the male clypeus is flat in profile. In
the majority of brachycistidines the
clypeus is convex and in Acanthetropis,
Brachycistis and Colocistis the clypeus
is concave in profile.

4. Maxillary palpal segments: normal (0),
reduced (1). The normal number of
maxillary palpal segments in tiphiids
is six. In Hadrocistis the maxillary palp
is polymorphic: 3 (slanskyae) or 5-
segmented (bicolor). In Brachycistina
the maxillary palp is 3-segmented.

5. Labial palpal segments normal (0),
reduced (1). The normal number of
labial palpal segments in tiphiids is
four. In Hadrocistis the labial palp is
polymorphic: 1-segmented in slanskyae
and 2-segmented in bicolor. In Brachy-
cistina the labial palp is 1-segmented.

6. Mandibular carina: absent (0), present
(1). In the primitive form of the
mandible, there are no external cari-
nae, angles or projections (as in
Fig. 11). In many brachycistidines the
mandible has a longitudinal ridge on
the outer surface extending from near

the base to a subapical position (as in
Fig. 2).

7. Frons: smooth above each antennal
socket (0), with arcuate carina and
apical warts (1). A peculiar modifica-
tion of the frons can be seen in
Brachycistis, Acanthetropis and Colocis-
tis (Figs 2, 4, 7). In these genera there is
a short crescentic carina above each
antennal socket. In most the carina
terminates on either end in a small
wart or tubercle. This is a feature
found only in this clade.

8. Hypostomal carina. Carina ending at
or outside mandibular socket (0),
ending inside mandibular socket (1).
In the majority of brachycistines and
in Tiphia the hypostomal carina termi-
nates midway on or outside of the
mandibular socket. The carina ends
inside the mandibular socket in Sedo-
maya, Brachycistis, Acanthetropis and
Colocistis.

9. Gular bridge: simple (0), with narrow
expansion over base of tongue (1).
Unlike the condition seen in tiphiines,
in some brachycistidines the gula has
a narrow expansion made up of
a broadened hypostomal carina that
overlaps the base of the tongue.

10. Pronotal dorsal surface: present (0),
absent (1). In Tiphia and Stilbopogon the
pronotum is dorsally expanded and
the dorsal edge is planar with the
scutum. In the rest of the Brachycisti-
dinae, with the exception of two
species of Colocistis, the pronotum
lacks a dorsal surface and is sub-
stantially shorter than the strongly
bulging scutum.

11. Stridulatory patch: absent (0), present
(1). Most tiphiids lack any kind of
stridulatory structure. However in the
majority of brachycistidine genera
males have a small stridulatory struc-
ture on the mesal surface of the
forecoxa, consisting of a defined,
slightly raised patch of dense fine
ridges (Fig. 27). This feature occurs in

Journal of Hymenoptera Research

both major brachycistidine clades with
a few notable, perhaps secondary,
absences in Brachycistina and Paraque-
mayn.

12. Scrobal sulcus: absent (0), present (1).
In brachycistidines a scrobal sulcus is
only present in Acanthetropis.

13. Hindwing discoidal veinlet: present
(0), absent (1). All of the brachycisti-
dine genera, except Acanthetropis and
Colocistis, have a short discoidal vein-
let present on the hindwing.

14. Propodeal transverse carina: absent
(0), present (1). A transverse carina
dividing the dorsal from the posterior
surfaces of the propodeum is seen
only in Acanthetropis, Colocistis and
Brachycistis.

15. Propodeum dorsomedially: convex
(0), longitudinally grooved (1). Anoth-
er modification of the propodeum is
the presence of a broad longitudinal
groove extending posteriorly from the
scutellar margin along the dorsal
surface of the propodeum. This feature
occurs in Colocistis and Acanthetropis.

16. Metapleural lobes: fused (0), digitate
(1) or broadly rounded (2). Tiphiines
have the metapleural lobes fused into
a platform between the midcoxae. In
the Brachycistidinae the metapleural
lobes are either small and slender or
flattened and apically rounded.

17. Hindcoxal dorsal carina: present (0),
absent (1). Throughout the Tiphiidae
the hindcoxa normally has a dorsal
carina that extends from the coxal base
towards the apex. This carina is absent
in Brachymaya, Hadrocistis and Brachy-
cistina.

18. Hindcoxal ventromedial carina: absent
(0), present (1). This is a unique feature
in Colocistis.

19. Basal petiolar carina: present (0), ab-
sent (1). In the Tiphiinae, and most
genera of Brachycistidinae the basal
metasomal tergum has a strongly ex-
panded lateral carina often completely
ringing the tergum, which may serve

to protect the petiolar socket. In
Stilbopogon this carina is absent and
the tergal base is simple where it
inserts into the petiolar socket.

20. Sternum I basal longitudinal carina
present (0), absent (1). Tiphiines have
a short longitudinal carina extending
down the middle of the basal gastral
sternum from the base of the plate. This
condition is also seen in Acanthetropis,
Brachycistelhts and Colocistis (Figs 28,
30). In the derived condition the first
gastral sternum is smooth basally with
no trace of a carina (Fig. 29).

21. Sternum II: simple (0), with medial
keel (1). The second gastral sternum is
generally unmodified, although Doli-
chetropis have a transverse submedial
carina. All species of Acanthetropis
have a longitudinal keel or fold ex-
tending longitudinally from the base.
In most Acanthetropis species this
structure consists of a sharp poster-
iorly hooked, keel-like carina in the
middle of this sternum. However, in
A. lamellatus this structure consists of
a longitudinal fold ending in a trans-
verse lamella.

22. Epipygium shape: rounded laterally
(0), laterally carinate or folded (1). The
epipygium, or apical metasomal ter-
gum, is generally narrowed and even-
ly rounded apically in the Tiphiidae.
In brachycistidines the epipygium can
be sharply folded or carinate laterally.

23. Digitus apex: broadly rounded (0),
apically elongate and acute (1). In
tiphiines the volsella is typically fist-
shaped, with the digitus and cuspis
forming short rounded apical lobes
(Figs 32, 35, 41, 46). However, in the
majority of brachycistidine genera the
digitus is greatly elongate and digitate,
or awl-shaped, extending apically (as
in Figs 36, 37).

24. Volsellar inner margin: smooth (0),
denticulate (1). Brachymaya, Hadrocis-
tis, Brachycistis, Dolichetropis, Paraqne-
maya and Sedomaya the inner margin of

Volume 15, Number 1, 2006

the volsella is denticulate (as in Figs 36,
38). This surface is smooth in the
remaining genera and in Tiphia.

RESULTS

Phylogenetic analyses. — Phylogenetic
analysis of the matrix yielded a single
optimal tree (Fig. 1A), with a tree length of
32, CI of 78 and RI of 83. Multistate
characters were treated as non-additive
and were equally weighted. Of these six
proved uninformative as they coded for
a single genus. Other characters used in the
key, such as the orientation of the hindw-
ing tCu vein and length of the forewing R
vein, were eliminated from consideration
early on because they only coded for
a single genus. Bootstrap values were

calculated after eliminating suboptimal
trees, resulting in the single tree in Fig. IB.
The values show strong to moderate
support for most of the branches. Stilbopo-
gon was found to be the basal taxon in the
subfamily in all analyses. The rest of the
genera fell into two well supported clades.
One clade consisting of Brachymaya, Ha-
drocistis and Brachycistina, based on the
lack of a dorsal coxal carina. The second
clade contains the remaining genera, based
on the presence of a mandibular carina and
a T-shaped carina at the base of metasomal
sternum I. The resulting generic relation-
ships are Stilbopogon + (Brachymaya +
(Hadrocistis + Brachycistina)) + (Paraquemaya
+ (Brachycistellus + (Dolichetropis + (Sedo-
maya + (Brachycistis + (Acanthetropis +
Colocistis)))))).

KEY TO MALES OF THE GENERA OF BRACHYCISTIDINAE

Forewing with two discoidal cells (as in Fig. 14) 2

Forewing with one discoidal cell (as in Fig. 15) 6

Sternum II with medial longitudinal ridge or keel extending posteriorly from base;
forewing R vein extending along costal margin more than two-thirds length of stigma
(Fig. 14) Acanthetropis Wasbauer

Sternum II without medial ridge; forewing R vein extending along costal margin for less
than two-thirds the length of stigma or not reaching costal margin at all (as in Figs 15,
17) 3

Mandible without longitudinal carina on external surface and with two apical teeth (Fig. 5);
maxillary and labial palpi highly reduced barely visible between mandibles, with three
and two or fewer palpomeres respectively Brachycistina Malloch

Mandible with longitudinal carina on external surface and with three apical teeth (as in
Fig. 4); maxillary and labial palpi clearly visible beneath head, extending as far as
occipital carina or further, with six and four palpomeres respectively 4

Maxillary palpi slender and short, barely reaching occipital carina; forewing R vein
extending along costal margin between one half and two-thirds stigmal length
(Fig. 20) Dolichetropis Wasbauer

Maxillary palpi robust and long, extending well beyond underside of head; forewing R
vein extending along costal margin less than one-third stigmal length or not even
reaching costal margin 5

Sternum I with short medial longitudinal carina extending posteriorly from base; forewing
tCu-1 arising at or before basal third of first submarginal cell (Fig. 19); digitus with
short, broadly rounded apex (Figs 39, 40) Colocistis Krombein

Sternum I smooth subbasally without medial carina; forewing tCu-1 arising beyond basal
third of first submarginal cell (Fig. 17); digitus with elongate, acute apex (Figs 36,
37) Brachycistis Fox

Journal of Hymenoptera Research

I Tiphia

•— Stilbopogon

1 9 22 23

>M>

1111

I Brachymaya

Hadrocistis
Brachycistina

-•-

5 19

I 1

Paraquemaya

15 23

Brachycistellus

-•-

2 0

-•-

Dolichetropis

Sedomaya

3 6 13 15

• •• >

2 112

IB r~Tiphia

12 14 23

1 1 o

Brachycistis

11 20

1 1
17 22

1 0

Acanthetropis
Colocistis

Stilbopogon

I Brachymaya

Hadrocistis

Brachycistina

Paraquemaya

Brachycistellus
— Dolichetropis
— Sedomaya

Brachycistis

Acanthetropis

Colocistis

Fig. 1. Cladograms of relationships among the ten genera of Brachycistidinae based on male characteristics.
Fig. 1A. Tree showing relationships among the genera. Numbers refer to characteristics discussed in the text.
Fig. IB. Tree showing Bootstrap values.

Hindwing cubitus forming a nearly straight line with transverse cubitus, meeting it at
more 130 angle or greater (Fig. 23); digitus short, blunt, without elongate apex (Figs 45,
46) Stilbopogon Mickel and Krombein

Hindwing cubitus arcuately curved, meeting transverse cubitus at less than 130 angle;
digitus usually with elongate, acute apex 7

Volume 15, Number 1, 2006

Acanlhelropis aequalis

Brachycistellus figitiformis

4. Brachycistis alcanor

5. Brachycistina acuta

6. Brachymaya mexicana

7. Colons Us crassa

czrf.

Dolichetropis flavida Hadrocistis slanskyae

1 0 . Paraquemaya pallida

12. Scdomaya glanuscnsis 13- Hadroasns slanskyae

1 1 . Quemaya marcida

Figs 2-13. Male head, with one or both antennae removed. Figs 2-12. Front view. Fig. 13. Ventral view.

Journal of Hymenoptera Research

14. Acantheiropis aequalis

1 6 . Brachycistina acuta

.8. Brachymaya mcxi carta

20. Dolichetropis flavida

Figs 14-21. Male fore and hindwings.

1 5 . Brachycislellus figitiformis

17. Brachycistis alcanor

19. Colocistis sfygia

2 1 . Hadrocistis slamkyae

Volume 15, Number 1, 2006

Paraquemaya bitincta

23. Quemaya marcida

Sedomaya glamisensis

Paraquemaya pallida

26.

Sedomaya glamiscnsis

Hrachycistis alcanor

28. 1/ \j, v 29.

Acanlhetropis lamellalus Dolichetropis jlavida

30. Colocistis stygia

Figs 22-24. Male fore and hindwings. Figs 25, 26. Lateral view of male head. Fig. 27. Ventral view of prothorax
and forecoxae. Figs 28-30. Ventral view of basal gastral sternum.

Palpi highly reduced, shorter than oral fossa, with fewer than five maxillary and three

labial palpomeres (Fig. 13) Hadrocistis Wasbauer

Palpi large, much longer than oral cavity; with six maxillary and four labial palpomeres 8

Propodeum heavily sculptured with transverse carina separating dorsal and posterior
surfaces; compound eyes and ocelli not enlarged, distance between midocellus and
nearest eye margin 4-5 MOD (Fig. 3) Brachycistellus Baker

Propodeum nearly smooth, without transverse carina; compound eyes and ocelli enlarged,
distance between midocellus and nearest eye margin not more than 3 MOD 9

Journal of Hymenoptera Research

31. 32.

Acanthetropis lamellatus

36. ^ 37.

Brachycistis alcanor

Brachycistellus figitiformis

35.
Brachycistina acuta

38. 39

Brachymaya mexicana

40
Colocistis stygia

Dolichetropis
flavidus

42.

Hadrocistis bicolor

Paraquemayabitincta *. Qumaya 46. gUfflup, Sedomaya

paupercula megalops glamisensis

Figs 31-47. Male genital capsule. Figs 31, 33, 34, 36, 38, 39, 42, 43, 45, 47. Ventral view. Figs 32, 35, 37, 40, 44, 46.
Lateral view.

Volume 15, Number 1, 2006

9 Mandible bidentate, without longitudinal carina on external surface (Fig. 6); oral carina

simple and gular area at base of mandible (traversed by oral carina) not dilated or

bulging Brachymaya Kimsey and Wasbauer

Mandible tridentate, with longitudinal carina on external surface; oral carina expanded
into tooth or lamella (Fig. 25) or gular area (traversed by oral carina) bulging near base
of mandible (Fig. 26) 10

10 Oral carina expanded into lamellate projection or tooth near mandibular base, gular area

not bulging near base of mandible (Fig. 25); forecoxa without stridulatory patch on

mesal surface Paraquemaya Kimsey and Wasbauer

Oral carina unmodified, gular area strongly bulging near base of mandible (Fig. 26);
forecoxa with stridulatory patch on mesal surface .... Sedomaya Kimsey and Wasbauer

CHECKLIST OF
THE BRACHYCISTIDINAE

Genus Acanthetropis Wasbauer

Figs 2, 14, 28, 31, 32
Acanthetropis Wasbauer 1958:139. Type species:
Acanthetropis lamellatus Wasbauer 1958:140.
Original designation. Distribution: USA:
South Dakota, south to Baja California Sur,
west to California.

Genus Brachycistina Malloch

Figs 5, 16, 34, 35
Brachycistina Malloch 1926:25. Type species
Brachycistis acuta Malloch 1926:25. Described
as a subgenus of Brachycistis Fox. Elevated
by Wasbauer (1966). Distribution: USA: Ar-
izona.

Species List

acuta Malloch 1926:25. {Brachycistina). Holotype

male; USA: Arizona, Arlington (NMNH).*

Species List

aequalis (Fox) 1899:284. (Brachycistis). Holotype
male; Nevada (ANSP).*

lamellatus Wasbauer 1958:140. (Acanthetropis).
Holotype male; Mexico: Baja California, La
Paz (CAS).*

noctivaga (Bradley) 1917:272. (Brachycistis). Ho-
lotype male; USA: Nevada, Las Vegas (CU).*

normalis (Malloch) 1917:8. (Brachycistis). Holo-
type male; USA: South Dakota, Cedar Pass
(NMNH).*

Genus Brachycistellus Baker

Figs 3, 15, 33
Brachycistellus Baker 1909:177. Type species:
Brachycistellus figitiformis Baker 1909:177.
Original designation. Distribution: USA:
southern California.

Species List
figitiformis (Baker) 1909:178. (Brachycistis). Holo-
type male; USA: California, Claremont

(NMNH).*

Genus Brachycistis Fox

Figs 4, 17, 27, 36, 37

Brachycistis Fox 1893:7. Type species: Brachycistis
petiolatus Fox 1893:8. Original designation.
Distribution: southwestern North America
(USA, northwestern Mexico).

Brachycistus (!) Cockerell and Casad. 1894:295.

Glyptomeopa Ashmead 1898:251. Type species:
Glyptometopa americana Ashmead 1898:251.
Monobasic. Synonymized by Wasbauer 1968.

Glytometopa (!) Fox 1899:289.

Brachycystis (!) Viereck 1906:289.

Bruesiella Mann 1914:182. Type species: Brue-
siella formicaria Mann 1914:182. Original des-
ignation. Synonymized by Kimsey 2006.

Eun/cros Mickel & Krombein 1942:666. Type
species: Eun/cros furtivus Mickel & Krombein
1942:667. Orig. designation. Synonymized by
Krombein 1967.

Aulacros Mickel & Krombein 1942:664. Type
species: Aulacros latior Mickel & Krombein
1942:665. Original designation. Synonymized
by Kimsey 2006.

Journal of Hymenoptera Research

Species List
agama (Dalla Torre) 1897:7. (Mutilla). Replace-
ment name for at rat a (Blake) 1879. Nee
Mutilla atratus Linnaeus 1767:966.
atrata (Blake) 1879:253. (Agama). Holotype
male; USA: Nevada (ANSP). Synonymized
by Wasbauer 1966.
contiguus Fox 1899:282. (Brachycistis). Syntype
males; USA: Nevada (ANSP). Synony-
mized by Wasbauer 1966.
nigritus Fox 1899:282. (Brachycistis). Syntype
males; USA: Nevada, Washington (ANSP).
Synonymized by Wasbauer 1966.
nudus Fox 1899:281. (Brachycistis). Holotype
male; USA: California (ANSP). Synony-
mized by Wasbauer 1966.
francisca (Mickel & Krombein) 1942:656.
(Glyptometopa). Holotype female; USA: Ca-
lifornia, San Francisco (CAS). Synony-
mized by Wasbauer 1968.
alcanor (Blake) 1879:264. (Agama). Holotype
male; USA: Arizona (ANSP).*
tysias Cameron 1900:396. Holotype male;
Mexico: northern Sonora (BMNH).* New
synonymy.
dubitatus Cameron 1900:397. (Brachycistis).
Holotype male; Mexico: northern Sonora
(BMNH).* New synonymy.
sciron Cameron 1900:400. (Brachycistis). Holo-
type male; Mexico: northern Sonora
(BMNH).* New synonymy.
cremastogaster Melander 1903:329. (Brachycis-
tis). Holotype male; Texas (Repository un-
known). Synonymized by Wasbauer 1966.
ampla (Blake) 1879:252. (Agama). Holotype male;

USA: Colorado (ANSP).
americana (Ashmead) 1898:251. (Glyptometopa).
Holotype female; USA: California, Alameda
Co. (NMNH).
arenivaga Bradley 1917:269. (Brachycistis). Holo-
type male; USA: California, Imperial Co. (CU).*
eriopis Bradley 1917:266. (Brachycistis). Holo-
type male: USA: California, Imperial Co.,
Calexico (CU). Synonymized by Wasbauer
1966.*
carinata Fox 1899:283. (Brachycistis). Holotype

male; USA: California (ANSP).
convergens (Mickel & Krombein) 1942:657. (Glyp-
tometopa). Holotype female; USA: California,
Santa Monica (NMNH).
convexa Wasbauer 1966:51. (Brachycistis). Holo-
type male; USA: California, Imperial Co.
(MCZ).

cruenta Wasbauer 1966:53. (Brachycistis). Holo-
type male; USA: Texas, Nueces Co., Corpus
Christi (CAS).*
curvata Malloch 1926:16. (Brachycistis). Holotype

male; USA: Arizona (NMNH).*
davidi Wasbauer 1966:56. (Brachycistis). Holo-
type male; USA: Texas, Jeff Davis Co. (CAS).
elegantula Cockerell & Casad 1894:295. (Brachy-
cistis). Holotype male; USA: New Mexico,
Dona Ana Co., Las Cruces (NMNH).*
exornatus Cameron 1900:402. (Brachycistis). Ho-
lotype male; Mexico: northern Sonora
(BMNH, type lost?).
formicaria (Mann) 1914:182. (Bruesiella). Holo-
type female; Mexico: Hidalgo (MCZ).
furtivus (Mickel & Krombein) 1942:667. (Eur-
ycros). Holotype female; USA: Arizona, Tuc-
son (UMSP).
glabrella Cresson 1865b:441. (Brachycistis). Holo-
type male; USA: New Mexico, Sandoval Co.
(ANSP).*

dentata Bradley 1917:263. (Brachycistis). Holo-
type male; USA: New Mexico, Jemez Spr.
(CU). Synonymized by Wasbauer 1966.
micrommata Bradley 1917:265. (Brachycistis).
Holotype male; USA: California, Alameda
Co. (CU). Synonymized by Wasbauer 1966.
utahensis Malloch 1926:17. (Brachycistis). Ho-
lotype male; USA: Utah (NMNH). Synon-
ymized by Wasbauer 1966.
dakotensis Malloch 1926:20. (Brachycistis).
Holotype male; USA: North Dakota, Edge-
mont (NMNH). Synonymized by Was-
bauer 1966.
melanocephala Malloch 1926:19. (Brachycistis).
Holotype male; USA: Utah, Uinta National.
Forest (NMNH). Synonymized by Was-
bauer 1966.
idiotes Cockerell 1895a:63. (Brachycistis). Holo-
type male; USA: New Mexico (Repository
unknown).
imitans Malloch 1926:23. (Brachycistis). Holotype
male; USA: California, Los Angeles Co.
(NMNH).*
inaequalis Fox 1899:282. (Brachycistis). Lectotype
male (design by Krombein 1954); USA:
California, Los Angeles Co. (NMNH).*
nevadensis Fox 1899:283. (Brachycistis). Holo-
type male; USA: Nevada (ANSP). Synony-
mized by Wasbauer 1966.*
nocticola Bradley 1917:289. (Brachycistis). Lap
sus calumni: name used in error in Expla-
nation of Figures, no holotype indicated or

Volume 15, Number 1, 2006

description given, as discussed by

Krombein 1954. Synonymized by Was-

bauer 1966.

indiscreta Fox 1899:284. (Brachycistis). Holotype

male; USA: Arizona, Cochise Co., Wilcox

(NMNH).*

stictinota Viereck 1906:190. {Brachycistis). Ho-
lotype male; USA: Kansas: Clark Co.
(SMEK). Synonymized by Wasbauer 1966.*
juncea Wasbauer 1966:35. (Brachycistis). Holo-
type male; USA: California, San Bernardino
(CAS).
lacustris Malloch 1926:24. {Brachycistis). Holo-
type male; USA: Nevada, Pyramid Lk.
(NMNH).*

hurdi Wasbauer 1966:39. {Brachycistis). Holo-
type male; USA: California, Riverside Co.
(CAS). Described as a subspecies of lacus-
tris Malloch. New synonymy.
latior (Mickel & Krombein) 1942:665. (Aulacros).
Holotype female; USA: California, Palm
Springs (CAS).
linsleyi Wasbauer 1966:21. {Brachycistis). Holo-
type male; Arizona: Globe (CAS).*
longula Wasbauer 1966:39. {Brachycistis). Holo-
type male; USA: California, Riverside Co.
(CAS).
nitida (Cresson) 1875:710. (Agama). Holotype
male; Colorado (ANSP).
lepidus (Blake) 1886:269. (Photopsis). Holotype
male; Colorado (ANSP). Synonymized by
Wasbauer 1966.*
nitida (Blake) 1886:267. {Photopsis). Holotype
male; Colorado (ANSP). Synonymized by
Wasbauer 1966.*
sobrina (Blake) 1886:268. {Photopsis). Holotype
male; Colorado (ANSP). Synonymized by
Wasbauer 1966.
petiolata Fox 1893:8. {Brachycistis). Holotype
male; Mexico: Baja California, Calmalli Mines
(CAS).*

gaudii Cockerell 1901:340. (Brachycistis). Lec-
totype male (designated herein); USA:
California, San Diego Co., La Jolla
(NMNH). Synonymized by Wasbauer 1966.
parva Malloch 1926:14. (Brachycistis). Holotype
male; USA: California (NMNH). Synony-
mized by Wasbauer 1966.
protracta Bradley 1917:270. (Brachycistis). Holo-
type male; USA: Arizona, Phoenix (ANSP).
semiatra Malloch 1926:24. (Brachycistis). Holo-
type male; USA: Washington, Adams Co.,
(NMNH)

subquadrata Fox 1899:282. (Brachycistis). Holo-
type male; USA: California, San Diego Co.,
San Diego (ANSP).

intermedia Malloch 1926:18. (Brachycistis). Ho-
lotype male; USA: California, San Diego
Co., San Diego (NMNH). Synonymized by
Wasbauer 1966.
texana Malloch 1926:12. (Brachycistis). Holotype

male; USA: Texas, Cotulla (NMNH).*
timberlakei Wasbauer 1966:70. (Bracliycistis). Ho-
lotype male; USA: California, San Diego Co.
(CAS).
triangularis Fox 1899:284. (Brachycistis). Holo-
type male; USA: Arizona (ANSP).*
verticalis Malloch 1926:19. (Brachycistis). Holo-
type male; USA: Arizona, Fort Grant
(NMNH).*
vigilax Cameron 1900:397. (Brachycistis).
Holotype male; Mexico: northern Sonora
(BMNH).

fulvilabris Cameron 1900:399. (Brachycistis).
Holotype male; Mexico: northern Sonora
(BMNH).* New synonymy.
arines Cameron 1900:401. (Brachycistis). Holo-
type male; Mexico: northern Sonora
(BMNH).* New synonymy.
ioachinensis Bradley 1917:267. (Brachycistis).
Holotype male; USA: California, Fresno
Co., Coalinga (CU).* New synonymy.
washingtona Malloch 1926:21. (Brachycistis).
Holotype male; USA: Washington, Ritzville
(NMNH). Synonymized by Wasbauer
1966.*
subcarinata Malloch 1926:11. (Brachycistis).
Holotype male; USA: Arizona, Higley
(NMNH). Synonymized by Wasbauer 1966.*
rugosa Malloch 1926:11. (Brachycistis). Holo-
type male; USA: Nevada, Golconda
(NMNH). Synonymized by Wasbauer
1966.*
uigrifrons Malloch 1926:16. (Brachycistis). Ho-
lotype male; USA: Arizona: Graham
Mount., Ash Crk. (NMNH). Synonymized
by Wasbauer 1966.*

Genus Br achy may a Kimsey & Wasbauer

Figs 6, 18, 38
Brachymaya Kimsey & Wasbauer 1998b:66. Type
species: Brachymaya mexicana Kimsey & Was-
bauer 1998:67. Original designation. Distri-
bution. USA: Baja California.

Species List
mexicana Kimsey & Wasbauer 1998b:67. (Brachy-
maya). Holotype male; Mexico: Baja Califor-
nia, San Augustine (LACM).

Genus Colocistis Krombein

Figs 7, 19, 30, 39, 40

Colocistis Krombein 1942:65. Type species: Colo-
cistis pilosa Krombein 1942:66. Original des-
ignation. Distribution: USA: California, Ar-
izona.

Aglyptacros Mickel & Krombein 1942:669. Type
species: Aglyptacros paxillatus (Mickel &
Krombein) 1942:673. Original designation.
Synonymized by Kimsey 2006

Species List

brevis (Fox) 1899:285. (Brachycistis). Holotype
male; USA: California (NMNH, lost?),*

castanea (Cresson) 1865a:388. (Brachycistis). Ho-
lotype male; USA: California (ANSP).*

crassa (Bradley) 1917:277. (Brachycistis). Holo-
type male; USA: California, Fresno Co.,
Coalinga (CU).*

eremi (Bradley) 1917:279. (Brachycistis). Holotype
male; USA: California, Imperial Co., Calexico
(CU).

paxillatus (Mickel & Krombein) 1942:673. (Aglyp-
tacros). Holotype female; USA: Colorado, La
Junta (NMNH).

pilosa Krombein 1942:66. (Colocistis). Holotype
male; USA: California, San Diego Co., San
Diego (NMNH).*

segredentatus (Mickel & Krombein) 1942:671.
(Aglyptacros). Holotype female; USA: Ari-
zona, Benson (MCZ).

sulcatus (Mickel & Krombein) 1942:674. (Aglyp-
tacros). Holotype female; USA: Texas, Valen-
tine (SMEK).
stygia (Bradley) 1917:276. (Brachycistis). Holo-
type male; USA: Arizona, Nogales (CU).*
thermarum (Bradley) 1917:274. (Brachycistis).
Holotype male; USA: Arizona, Hot Springs
(CU).

Genus Dolichetropis Wasbaue

Figs 8, 20, 29, 41
Dolichetropis Wasbauer 1968a:191. Type species:
Dolichetropis flavida Wasbauer 1968a:192.

Journal of Hymenoptera Research

Original designation. Distribution: Mexico:
Baja California Sur.

Species List
flavida Wasbauer 1968a:192. (Dolichetropis). Ho-
lotype male; Mexico: Baja California Sur, San
Ignacio (CAS).*

Genus Glyptacros Mickel & Krombein

Glyptacros Mickel & Krombein 1942:660. Type
species: Glyptacros angustior Mickel & Krom-
bein 1942:660. Original designation. Distribu-
tion. Southwestern USA: California, Arizona,
Utah, Wyoming, New Mexico, Texas; Mexico:
Baja California Sur, Chihuahua, Hidalgo,
Sonora.

Xeroglypta Mickel & Krombein 1942:663. Type
species: Xeroglypta egregia Mickel & Krombein
1942:663. Original designation. Synonymized
by Kimsey 2006.

Astigmometopa Mickel & Krombein 1942:668.
Type species: Astigmometopa emarginata
Mickel & Krombein 1942:668. Original desig-
nation. Synonymized under Brachycistis by
Wasbauer 1971. Removed from synonymy
under Brachycistis and synonymized under
Glyptacros by Kimsey 2006.

Species List

angustior Mickel & Krombein 1942:660. (Glypta-
cros). Holotype female; USA: Arizona
(ANSP).

ashmeadi Mickel & Krombein 1942:661. (Glypta-
cros). Holotype female; no locality given
(NMNH).

egregia (Mickel & Krombein) 1942:663. (Xero-
glypta). Holotype female; USA: California,
Palm Springs (UMSP).

emarginata (Mickel & Krombein) 1942:668. (As-
tigmometopa). Holotype female; USA: Texas,
Valentine (SMEK). "

eureka (Banks) 1912:202. (Glyptometopa). Holo-
type female; USA: Arizona, Palmerlee (MCZ).

Genus Hadrocistis Wasbauer

Figs 9, 13, 21, 42
Hadrocistis Wasbauer 1968a: 185. Type species:
Hadrocistis bicolor Wasbauer 1968a:187. Orig-
inal designation. Distribution: USA: Califor-
nia, Imperial Co.; Mexico: Baja California,
Sinaloa.

Volume 15, Number 1, 2006

Species List
bicolor Wasbauer 1968a:187. (Hadrocistis). Holo-

type male; USA: California, Imperial Co.,

6 mi w Glamis (CAS).*
slanskyae Wasbauer 1968a:188. (Hadrocistis). Ho-

lotype male; USA: California, Imperial Co.,

6 mi w Glamis (CAS).*

Genus Paraquemaya Kimsey & Wasbauer

Figs 10, 22, 25, 43, 44
Paraquemaya Kimsey and Wasbauer 1998b:67.
Type species: Paraquemaya pallida Kimsey &
Wasbauer 1998:71. Distribution. USA: Ari-
zona; Mexico: Baja California Sur, Sonora.

Species List

bajaensis Kimsey and Wasbauer 1998b:69. (Para-
quemaya). Holotype male; Mexico: Baja Cali-
fornia Sur, 14 mi s El Arco (UCRC).*

bitincta Kimsey and Wasbauer 1998b:70. (Para-
quemaya). Holotype male; USA: Arizona,
Pima Co., Organ Pipe (UCRC).*

maxima Kimsey and Wasbauer 1998b:71. (Para-
quemaya). Holotype male; Mexico: Sonora,
22 km se Quitovac (UCRC).*

pallida Kimsey and Wasbauer 1998b:71. (Para-
quemaya). Holotype male; Mexico: Baja Cali-
fornia Sur, 4 mi wsw Miraflores (UCDC).*

Genus Sedomaya Kimsey & Wasbauer

Figs 12, 24, 26, 47
Sedomaya Kimsey & Wasbauer 1998b:72. Type
species: Sedomaya glamisensis Kimsey & Was-
bauer. Original designation. Distribution.
USA: California, Imperial Co.

Species List
glamisensis Kimsey & Wasbauer 1998b:72. (Se-
domaya). Holotype male; USA: California,
Imperial Co., Glamis (UCDC).

Genus Stilbopogon Mickel & Krombein

Figs 11, 23, 45, 46
Stilbopogon Mickel & Krombein 1942:658. Type
species: Stilbopogon alutacea Mickel & Krom-
bein 1942:659. Original designation. Synony-
mized by Wasbauer 1971 under Brachycistis
Fox. Distribution. USA: California, Nevada,

Arizona, New Mexico, Texas; Mexico: Baja
California, Sonora; Costa Rica: Guanacaste;
Colombia: Magdalena.
Quemaya Pate 1947:139. Type species: Brachycis-
tis paupercula Bradley 1917:282. Original
designation. Synonymized by Kimsey 2006.

Species List

alutacea Mickel & Krombein 1942:659. (Stilbopo-
gon). Holotype female; USA: New Mexico,
Deming (NEW YORK).

arenicola (Wasbauer) 1967:169. (Quemaya). Holo-
type male; USA: California, Imperial Co.,
6 mi nw Glamis (CAS).*

confusa (Kimsey & Wasbauer) 1998b:42. (Que-
maya). Holotype male; Mexico: Sonora,
6 km nnw San Carlos (UCDC).*

costaricana (Kimsey & Wasbauer) 1998b:43.
(Quemaya). Holotype male; Costa Rica: Gua-
nacaste Prov., 14 km s Canas (EMUS).*

eurekensis (Kimsey & Wasbauer) 1998b:43. (Que-
maya). Holotype male; USA: California, Inyo
Co., Eureka Valley, dunes (UCDC).*

inermis (Malloch) 1924:23. (Brachycistis). Holo-
type male; USA: Arizona: Higley (NMNH).*

marcida (Bradley) 1917:283 (Brachycistis). Holo-
type male; USA: California, Imperial Co.
(CU).*

megalops (Kimsey & Wasbauer) 1998b:44. (Que-
maya). Holotype male; USA: California, Riv-
erside Co., 5 mi nw Desert Center (UCDC).*

mexicana (Kimsey & Wasbauer) 1998b:43. (Que-
maya). Holotype male; Mexico: Baja Califor-
nia, 1 km s El Rosario (UCDC).*

paupercula (Bradley) 1917:282. (Brachycistis). Ho-
lotype male; USA: California, Imperial Co.
(CU).*

perpunctata (Cockerell) 1895b:291. (Brachycistis).
Holotype male; USA: New Mexico, Dona Ana
Co., Las Cruces (Repository unknown).

sonorensis (Kimsey & Wasbauer) 1998b:46. (Que-
maya). Holotype male; Mexico: Sonora,
6 km nnw San Carlos (UCDC).*

ACKNOWLEDGMENTS

Our thanks go to the collections and collection
managers who made this study possible, and to Maria
Garcia who rendered many of the ink drawings.

LITERATURE CITED

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Psyche 8: 251.

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from California. Invertebrata Pacifica 1: 177-178.

Banks, N. 1912. New Scolioidea. Canadian Entomologist
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Blake, C. A. 1879. Catalog of the Mutillidae of North
America with descriptions of new species. Trans-
actions of the American Entomological Society 7:
243-254.

Blake, C. A. 1886. Monograph of the Mutillidae of
North America. Transactions of the American
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Bradley, J. C. 1917. Contributions toward a monograph
of the Mutillidae and their allies of America north
of Mexico. IV. A review of the Myrmosidae.
Transactions of the American Entomological Society
43: 247-290.

Brothers, D. J. 1975. Phylogeny and classification of
the aculeate Hymenoptera, with special reference
to Mutillidae. University of Kansas Science Bulletin
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Brothers, D. J. and J. M. Carpenter. 1993. Phylogeny of
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Hi/menoptera Research 2: 227-301.

Cameron, P. 1900. Fossores. Biologia Centrali-Americana
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Cockerell, T. D. A. 1895a. Descriptions of new
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Cockerell, T. D. A. 1895b. Descriptions of new
Hymenoptera. Transactions of the American Ento-
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Cockerell, T. D. A. 1901. Species of Brachycistis from
southern California. Canadian Entomologist 45:
340-341.

Cockerell, T. D. A. and J. E. Casad. 1894. New species
of Mutillidae. Entomological News, Philadelphia 5:
293-296.

Cresson, E. T. 1865a. Descriptions of some new species
of Mutilla from California. Proceedings of the
Entomological Society of Philadelphia 4: 385-390.

Cresson, E. T. 1865b. Catalog of Hymenoptera in the
collection of the Entomological Society of Phila-
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the Entomological Society of Philadelphia 4: 426^488.

Cresson, E. T. 1875. Report upon the collections of
Hymenoptera. Chapter VII in: Wheeler, W. M. ed.
U.S. Geographic surveys west of the 100"' meridian. V.
Zoology.

Dalla Torre, C. G. 1897. Catalogus Hymenopterorum, 1.
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Farris, J. S. 1988. Hennig86, version 1.5 manual/
software and MSDOS program. Distributed by
the author. Port Jefferson Station, New York.

Fox, W. J. 1893. Report on some Mexican Hymenop-
tera, principally from Lower California. Proceed-
ings of the California Academy of Sciences (2)4: 1-25.

Fox, W. J. 1899. The North American Mutillidae.
Transactions of the American Entomological Society
25: 219-300. '

Kimsey, L. S. 1991. Relationships among the tiphiid
wasp subfamilies. Systematic Entomology 16:
427-438.

Kimsey, L. S. 2006. Competing taxonomies: reexami-
nation of the female-based genera of Brachycisti-
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the American tiphiid genus Quemaya Pate, journal
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Kimsey, L. S. and M. S. Wasbauer. 1998b. New Genera
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Malloch, J. R. 1924. A new species of the genus
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Melander, A. L. 1903. Notes on the North American
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Volume 15, Number 1, 2006 25

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J. HYM. RES.
Vol. 15(1), 2006, pp. 26-37

Fauna and Biogeography of the Bees and Wasps of the Cook Islands

(Hymenoptera Aculeata)

Michael Kuhlmann

Institute of Landscape Ecology, University of Miinster, Robert-Koch-Str. 26, D-48149 Munster,

Germany; email: kuhlmmi@uni-muenster.de

Abstract— A total of 19 species of bees and aculeate wasps is currently known from the Cook
Islands with six of them recorded for the first time: Pison spec. 2, Sceliphron laetum (F. Smith, 1856),
Ceratina (Neoceratina) dentipes Friese, 1914, Megachile {Eutricharaea) spec. 1, M. (E.) spec. 2. The
following three species are presumably indigenous: Anterhynchium (Epiodynerus) rufipes (Fabricius,
1775), Parodynerus bicinctus (Fabricius, 1781), Tachysphex f.fanuiensis Cheesman, 1928. Three more
species are possibly indigenous but their status remains uncertain due to taxonomic problems:
Pison spec. 1, Pison spec. 2, Megachile (Eutricharaea) spec. 1. A comparative analysis of faunistic data
from other Pacific islands showed that the indigenous aculeate fauna of the Cook Islands is
extremely depauperate. In general the number of species, especially parasitic taxa, and endemics
decline from west to east with the remote archipelagos of Polynesia possessing the most
depauperate aculeate faunas consisting only of widespread species.

Traditionally the oceanic Pacific islands perate. However, isolation and a highly
are divided into Micronesia, Melanesia and diverse environment can lead to remark-
Polynesia, originally on the basis of their able radiations, such as the 60 species of
indigenous aboriginal peoples, with the Hylaeus (Nesoprosopis) (Colletidae) and 99
Cook Islands as a part of the latter Odynerus species (Vespidae) in the Hawai-
(Crocombe 2001). Polynesia is a vast tri- ian Islands (Nishida 1994, Daly and Mag-
angular area with the edges formed by nacca 2003) and the endemic genus Ech-
Hawaii in the north, Easter Island (Rapa- thralictus in Samoa (Michener 2000). In-
nui) in the southeast and New Zealand in terestingly, endemic forms among the bees
the southwest, including the territories of and wasps seem to have developed only on
Hawaii, Kiribati (part), Tuvalu, Tokelau, the high islands. The low coral atolls have
Wallis & Futuna, Samoa, American Samoa, only a sparse, introduced fauna derived
Tonga, Niue, Cook Islands, French Poly- mainly from the Oriental region (Krombein
nesia, Pitcairn, Easter Island and New 1949b, 1950). Except for Hawaii (Nishida
Zealand (Crocombe 2001). Other than 1994, Daly & Magnacca 2003, Snelling
New Zealand, that has a continental origin, 2003) and Samoa (Perkins and Cheesman
the islands of Polynesia are of volcanic 1928, Williams 1928) the bees and aculeate
origin. To colonize these isolated islands wasps of Polynesia are poorly studied
the terrestrial fauna and flora had to cross (compilation in Williams 1947). It is the
large areas of open ocean. Because only intention of his study to present the results
a small fraction of the bee and wasp of an investigation of the bees and aculeate
species is capable of long distance dispers- wasps of the islands of Rarotonga and
al (Michener 1979, 2000) the fauna derived Mangaia made in 2004, including other
from relatively few immigrants. As a result available data about the Cook Islands. The
oceanic island faunas are usually depau- biogeographic situation is discussed in

Volume 15, Number 1, 2006

a Polynesian and Pacific context including
a summary of data available for Pacific
Islands.

MATERIALS AND METHODS

The Cook Islands consist of 15 islands
located in the central South Pacific between
the Society Islands to the east and Tonga
and Samoa to the west. During a stay on
the Cook Islands from 13th September to 7lh
October 2004 I collected on the two largest
and most southerly of its 15 islands:
Rarotonga with the capital Avarua
(67 km2) and Mangaia (51 km2). Both have
a tropical climate with Rarotonga being
a geologically young (about 2 million
years) high volcanic island (maximum
elevation 653 m). The geologically old
(about 18 million years) Mangaia is classi-
fied as a makatea island consisting of an
uplifted karstified limestone rim encircling
a central volcanic core (maximum eleva-
tion 169 m) (Hein et al. 1997). The speci-
mens collected there are deposited in my
collection. Reference material is located in
the collections of various specialists (see
acknowledgements). Additional records
were taken from the Cook Islands Bio-
diversity Database (McCormack 2004). In
the faunistic part families are in systematic,
species in alphabetical order.

LIST OF WASPS AND BEES OF THE
COOK ISLANDS

Chrysididae

Chrysis sp.

Distribution. — This unidentified species
is known from Rarotonga (Totokoitu Re-
search Station) and Atiu (McCormack
2004). Probably introduced.

Biology. — Cuckoo wasps of the genus
Chrysis are known as parasites of a broad
spectrum of aculeate wasps and bees
(Kimsey 1990). The host of this species is
unknown.

Specimens examined. —

Vespidae

Anterhynchium (Epiodynerus) rufipes (Fabri-
cius, 1775)

Distribution. — Widespread in the Pacific
region: Australia, New Guinea, Fiji, Tonga,
Samoa, Tuvalu, Tokelau, Niue, Society
Islands, Marquesas (Giordani Soika 1957,
Carpenter, pers. comm.). Probably indige-
nous to the Cook Islands.

Previous records are from Rarotonga
(1925, leg. G. Wilder; 1977, leg. Kraus,
1999), Atiu (1925, leg. G. Wilder; 1976, leg.
Krauss) and Aitutaki (1960, 1976, leg.
Krauss; Krauss 1961) but missing on
Pukapuka (McCormack 2004). Probably
present on most islands of the archipelago.

Biology. — The species can be regularly
observed hunting or visiting flowers in
gardens, along road sides and on fallow
land. Two males from Mangaia carry some
phoretic mites.

Specimens examined. — Rarotonga: Arorangi,
fallow land, 10 m (21 12'54"S 159 49'42"W),

2 9, 19. ix. 2004; Arorangi, upper beach, 5 m
(21 12'45"S 159 49'39"W), 1 ;, 21.ix.2004; 1 9,
4.X.2004; Mangaia: Ivirua, on Makatea, 30 m
(21 55'15"S 157°53'21"W), 1 9, 1 J, 22.ix.2004; 3 9,

3 $, 27.ix.2004.

Delta esuriens okinawae Giordani Soika, 1986

Distribution. — The specimens correspond
best to this subspecies that is known from
China and Taiwan. It differs very little
from typical D. esuriens from India and
might be conspecific (Carpenter, pers.
comm.). Introduced to the Cook Islands.

Previously known only from Rarotonga
(Tupapa, xii.2002, leg. G. McCormack)
(McCormack 2004).

Biology. — Species of Delta construct free
clay nests on rocks, houses or wood and
provision the cells with paralyzed cater-
pillars (Mader 2000).

Specimens examined. — Rarotonga: Arorangi,
upper beach, 5 m (21 14'20"S 159 49'40"W),
1 o\ 18.ix.2004; Mangaia: S Ivirua, on Makatea,
30 m (21 55'52"S 157 52'42"W), 1 9, 26.ix.2004.

Pachodynerus nasidens (Latreille, 1817)

Distribution. — This species originated
from the Neotropics and is now wide-
spread throughout the Pacific region (Car-
penter, pers. comm.). Introduced to the
Cook Islands.

In the Cook Islands this species is very
common and known from Rarotonga, Atiu,
Aitutaki and Pukapuka (Krauss 1961,
McCormack 2004). P. nasidens is probably
present on most islands of the archipelago.

Biology. — The species can be regularly
observed hunting or visiting flowers in
gardens, along road sides and on fallow
land, with a preference for the introduced
Mexican Fire-plant (Euphorbia cyathovhora
J. A. Murray) (Euphorbiaceae).

Specimens examined. — Rarotonga: Arorangi,
upper beach, 5 m (21 14'20"S 159 49'40"W),
1 9, 5 $, 15.ix.2004; 2 3, 18.ix.2004; Arorangi,
fallow land, 10 m (21 12'54"S 159 49'42"W), 2 9,
5 3, 19.ix.2004; Arorangi, upper beach, 5 m
(21 12'45"S 159 49'39"W), 1 9, 7$, 20.ix.2004; 2 3,
21.ix.2004; 1 3, 4.X.2004; Mangaia: Ivirua, on
Makatea, 30 m (21 55'15"S 157 53'21"W), 2 $,
22. ix. 2004; Ivirua, Taro gardens, 30 m
(21 55'20"S 157 53'53"W), 1 9, 1 3, 23.ix.2004; S
Ivirua, on Makatea, 30 m (21 55'52"S
157°52'42"W), 1 3, 26.ix.2004.

Parodynems bicinctns (Fabricius, 1781)

Distribution. — Widespread in the Pacific
region: New Guinea, Philippines, Fiji,
Tonga, Tuvalu, Samoa, Cook Islands, Soci-
ety Islands, Gambier Islands, Marquesas,
Tuamotu Archipelago (Giordani Soika
1957, Carpenter, pers. comm.). Probably
indigenous to the Cook Islands.

In the Cook Islands this species is very
common and known from Rarotonga
(1979, leg. Krauss), Atiu (1976, leg. Krauss),
Mitiaro (1979, leg. ?Krauss) and Aitutaki
(1977, leg. Krauss; Krauss 1961) but miss-
ing on Pukapuka (McCormack 2004). Prob-
ably present on most islands of the
archipelago.

Biology. — This very common species can
be regularly observed hunting or visiting

Journal of Hymenoptera Research

flowers in gardens, along road sides
and on fallow land, with preference
for he introduced Mexican Fire-plant (Eu-
phorbia cyathophora J.A. Murray) (Euphor-
biaceae).

Specimens examined. — Rarotonga: Arorangi,
upper beach, 5 m (21 14'20"S 15949'40"W),
1 9, 15.ix.2004; 1 3, 18.ix.2004; Arorangi, fallow
land, 10 m (21 12'54"S 159 49'42"W), 1 9,
19. ix. 2004; Arorangi, upper beach, 5 m
(21T2'45"S 159 49'39"W), 1 9, 73, 20.ix.2004;
1 9, 6 3, 21.ix.2004; 1 9, 2 3, 4.X.2004; Mangaia:
Ivirua, Taro gardens, 30 m (21 55'20"S
157 53'53"W), 1 9, 1 3, 23.ix.2004; S Ivirua, on
Makatea, 30 m (21 55'52"S 157 52'42"W), 1 9,
26.ix.2004; Ivirua, on Makatea, 30 m (21 55'15"S
157 53'21"W), 2 9, 5 3, 22.ix.2004; 2 9, 1 3,
27.ix.2004.

Polistes jokahmnae Radoszkowski, 1887

Distribution. — India, China, Mongolia,
Korea, Japan, Taiwan, Society Islands,
Tuamotu Archipelago; introduced to Ha-
waii (Carpenter 1996, as P. jadwigae).
Probably introduced to the Cook Islands.

Previous records are from Rarotonga
(iv.1925, leg. G. Wilder; xii.1977, Titika-
veka, lowlands, leg. N.L.H. Krauss;
xii.1977, Avarua lowlands, leg. N.L.H.
Krauss, v. 2002, Avana valley, leg. G.
McCormack), Atiu (xi.1977, central plateau,
leg. N.L.H. Krauss) and Aitutaki (ii.1960,
leg. N.L.H. Krauss; xi.1977, Tautu, leg.
N.L.H. Krauss; Krauss 1961) but missing
on Pukapuka (McCormack 2004).

Biology. — This social paper wasp is ubi-
qitous in open habitats on the Cook Islands
but less common than P. olivaceus. The
species builds free hanging nests in bushes
and trees.

Specimens examined. — Rarotonga: Arorangi,
Raemaru trail, 50 m (21 14'06"S 159 49'19"W),
3 9/ 17.ix.2004; Arorangi, upper beach, 5 m
(21 12'45"S 159 49'39"W), 1 9, 20.ix.2004; Man-
gaia: Ivirua, on Makatea, 30 m (21 55'15"S
157 53'21"W), 3 9, 22.ix.2004; 4 9, 27.ix.2004;
S Ivirua, on Makatea, 30 m (21 55'52"S
157 52'42"W), 1 9, 26.ix.2004.

Volume 15, Number 1, 2006

Polistes olivaceus (de Geer, 1773)

Distribution. — East Africa, South Asia,
Australia and common on archipelagos of
the Indian and Pacific ocean (Carpenter
1996). Probably introduced to the Cook
Islands.

Very common in the Cook Islands but
missing in Tongareva (= Penrhyn) and
Pukapuka (Krauss 1961, McCormack 2004).

Biology. — This social paper wasp is ubi-
qitous in open habitats on the Cook Islands
and builds free hanging nests in bushes
and trees.

Specimens examined. — Rarotonga: Arorangi,
upper beach, 5 m (21 14'20"S 159 49'40"W), 7
9, 15.ix.2004; 1 9, 18.ix.2004; Arorangi, Raemaru
trail, 50 m (21 14'06"S 15949'19"W), 1 9,
17. ix. 2004; Arorangi, fallow land, 10 m
(2112'54"S 159 49'42"W), 2 9, 19.ix.2004; Aror-
angi, upper beach, 5 m (21 12'45"S
159 49'39"W), 1 9, 20.ix.2004; 2 9, 4.X.2004;
Mangaia: Ivirua, on Makatea, 30 m (21 55'15"S
157 53'21"W), 1 9, 22. ix. 2004; S Ivirua, on
Makatea, 30 m (21 55'52"S 157 52'42"W), 1 9,
26.ix.2004.

Ampulicidae
Ampulex compressa (Fabricius, 1781)

Distribution. — Ethiopia, Kenya, Tanza-
nia, Arabian Peninsula, Madagascar, Sey-
chelles, Reunion, Mauritius, India, Sri
Lanka, Bangladesh, Singapore, south Chi-
na, Indonesia, Philippines, Australia, New
Caledonia (Pulawski 2003). To help control
the cockroach Peripilaneta americana (Lin-
naeus, 1758) between 1955 and 1958 its
parasitoid A. compressa was introduced to
Rarotonga from Hawaii, where it had been
introduced from New Caledonia in 1941
(Walker & Deitz 1979, McCormack 2004).
The conspicuous sphecid has not been
observed for many years and might be
extinct.

Biology. — All Ampulex species hunt cock-
roaches and nest in preexisting cavities
(Bohart & Menke 1976).

Specimens examined. —

Sphecidae
Sceliphron caementarium (Drury, 1773)

Distribution. — The origin of this species
is North and Central America, now rang-
ing worldwide (Pulawski 2003). Intro-
duced to the Cook Islands. Common on
Rarotonga (McCormack 2004) and also
present on Aitutaki (Krauss 1961).

Biology. — Species of Sceliphron construct
free clay nests on rocks, houses or wood and
provision the cells with spiders (Bohart and
Menke 1976). This synanthropic species is
common on Rarotonga where it can be
frequently observed flying along walls and
roofs even in the centre of the capital Avarua.

Specimens examined. — Rarotonga: Arorangi,
fallow land, 10 m (21°12'54"S 159 49'42"W),
8 6*/ 19.ix.2004; Arorangi, upper beach, 5 m
(21T2'45"S 159°49'39"W), 1 J, 20.ix.2004.

Sceliphron laetum (F. Smith, 1856)

Distribution. — India, Indonesia, New
Guinea, Australia, New Zealand, Guam,
Mariana Islands (Pulawski 2003). First re-
cord for the Cook Islands of this introduced
species. Like other species of Sceliphron S.
laetum is easily introduced to islands by
ships. According to Krombein (1949b), this
species accompanied a ship that sailed over
3,000 km from the Solomons to Guam in
1945 and established a colony there.

Biology. — Species of Sceliphron construct
free clay nests on rocks, houses or wood
and provision the cells with spiders (Bo-
hart and Menke 1976). This species is less
common on Rarotonga than S. caementar-
ium and only known from one locality.

Specimens examined. — Rarotonga: Arorangi,
fallow land, 10 m (21 12'54"S 159 49'42"W), 2
9, 2 $, 19.ix.2004.

Crabronidae
Pison sp. 1

Distribution. — About half of the 196
worldwide known species of the genus
Pison occur in Australia and the Pacific

Journal of Hymenoptera Research

region with 26 of them known from
Oceania (Pulawski 2003). The genus is
badly in need of revision before the two
species found on the Cook Islands can be
identified. McCormack (2004) listed this
bigger species as P. tahitense (de Saussure
1867) but it differs from it in the following
points: mesopleuron sparsely punctate,
with punctures several diameters apart
(subcontiguous in P. tahitense), the propo-
deal posterior surface is punctate (trans-
versely rugulosopunctate in P. tahitense)
and the wings are translucent (moderately
infumate in P. tahitense) (Pulawski pers.
comm.). Krauss (1961) mentions P. hospes
(F. Smith 1879) from Aitutaki. This species
is known from Singapore, Philippines,
New Guinea, Micronesia, Hawaii, Fiji,
Tonga, Samoa, Marquesas and Society
Islands (Yasumatsu 1953, Pulawski 2003).
The two species I collected on Rarotonga
and Mangaia, about 250 km south of
Aitutaki, are clearly different from P. liospes
(Pulawski pers. comm.). Due to the difficult
taxonomic situation in this genus the
identity of P. hospes reported by Krauss
(1961) needs to be confirmed. Beside P.
hospes and P. tahitense three more Pison
species are known to occur in regions
adjacent to the Cook Islands: P. iridipenne
F. Smith, 1879 (Australia, New Guinea,
Philippines, Hawaii, Micronesia, Fiji, Sa-
moa, Society Islands, Tuamotu Archipelago,
Marquesas), P. ignavwn R. Turner, 1908
(Australia, New Guinea, Philippines, Tai-
wan, Micronesia, New Caledonia, Fiji,
Samoa, Society Islands, Marquesas) and P.
iinpitnctatinn R. Turner, 1912 (New Guinea,
Society Islands, Marquesas) (Yasumatsu
1953, Pulawski 2003). The latter species
was not available for comparison and the
former two are different from the species
collected (Pulawski pers. comm.). Both
species collected during the survey are
probably indigenous to the Cook Islands.

Previously known from Rarotonga (1999;
iii.2003, leg. McCormack), Mangaia and
Pukapuka (ii.2004, leg. McCormack)
(McCormack 2004).

Biology.— Species of Pison construct ei-
ther free clay cells or partitions holes in
timber and provision the cells with spiders
(Krombein 1949b).

Specimens examined.— Rarotonga: Arorangi,
Raemaru trail, 50 m (2L14'06"S 159 49'19"W),
1 9, 17.ix.2004; Mangaia: S Ivirua, on Makatea,
30 m (21 55'52"S 157 52'42"W), 15 9, 26.ix.2004;
Ivirua, on Makatea, 30 m (21 55'15"S
157 53'21"W), 6 9, 22.ix.2004; 1 9, 27.ix.2004.

Pison sp. 2

Distribution. — For general comments on
the genus see previous species. First record
of this smaller species for the Cook Islands.
Probably indigenous.

Biology. — See previous species.

Specimens examined. — Rarotonga: Arorangi,
Raemaru trail, 50 m (21 14'06"S 159 49'19"W),
3 9, 3 j\ 17.ix.2004; Arorangi, upper beach, 5 m
(21 12'45"S 159 49'39"W), 1 9, 20.ix.2004; 4 9,
21.ix.2004.

Tachysphex fanuiensis fanuiensis Cheesman,
1928

Distribution. — New Caledonia, Loyalty
Islands, Society Islands, Tuamotu Archi-
pelago, Marquesas. The subspecies T. f.
howeanns Pulawski, 1977 is only known
from Lord Howe Island and T. f. corallimts
Pulwaski, 1977 occurs on the north
and east coast of Australia, New Guinea
and the Solomon Islands (Pulawski 1977).
A female was collected in 1925 on a ship
about 750 km SE of Pitcairn flying
over a bulwark (Pulawski 1977). Obviously
T. /. fanuiensis can be easily introduced to
other islands. Indigenous to the Cook
Islands.

Earlier records are from Rarotonga (1977,
1979, leg. Krauss; ii.2003, leg. G. McCor-
mack) and Mauke (1976, leg. Krauss)
(McCormack 2004).

Biology. — This species is very common
on sandy and rocky parts of the upper
beach where it nests in the ground or in
existing cavities of rocks. It can also be

Volume 15, Number 1, 2006

frequently found in sparsely vegetated
places further inland. Females prey upon
cockroaches which sit exposed on leaves
(Pulawski 1977). Males often visit the
introduced Mexican Fire-Plant (Euphorbia
cyathophora J. A. Murray) (Euphorbiaceae)
for nectar.

Specimens examined. — Rarotonga: Arorangi,
upper beach, 5 m (21 14'20"S 159 49'40"W), 8
9, 12 3, 15.ix.2004; Arorangi, Raemaru trail,
50 m (21°14'06"S 159 49'19"W), 2 3, 17.ix.2004;
Arorangi, upper beach, 5 m (21 12'45"S
15949'39"W), 3 9, 7 $, 20. ix. 2004; 1 6\
21.ix.2004; 1 9, 2 $, 4.X.2004; Mangaia: Ivirua,
Taro gardens, 30 m (21 55'20"S 157 53'53"W), 1
9, 10 6\ 23.ix.2004.

Megachilidae

Lithurgus (Litlwrgus) scabrosus
(F. Smith, 1859)

Distribution. — India, Indonesia, New
Guinea and many Pacific Islands east-
wards up to Tahiti (Michener 1965, Snel-
ling 2003). Due to their nesting in wood
these bees have an excellent ability to cross
water barriers, e.g. on ships (Michener
1965). Probably introduced to the Cook
Islands by ancient Polynesians.

Previous records are from Rarotonga
(iii.1979, leg. N.L.H. Krauss) and Aitutaki
(1977, leg. N.L.H. Krauss) (McCormack2004).

Biology. — Six of the seven collected speci-
mens carry the phoretic mite Chaetodactylus
ludwigi (Trouessart, 1904), some of them in
large amounts. This mite is known from
Lithurgus species from Madagascar, India,
Java, New Caledonia, Moorea Island near
Tahiti and Eastern Caroline Islands (Fain
and Pauly 2001). I found L. scabrosus
nesting in decaying wooden poles and like
Pauly & Munzinger (2003) I observed
a female visiting an Ipomoea species -
Coastal Morning-Glory (Ipomoea littoralis
Blume) (Convolvulaceae). Species of this
subgenus are known to be oligoleges of
Malvaceae (such as the widely distributed
Hibiscus) (Snelling 2003) and Convolvula-
ceae (such as Ipomoea pes-caprae (L.) Sweet,

a widely distributed plant on pantropical
seashores) (Pauly et al. 2001).

Specimens examined. — Rarotonga: Arorangi,
upper beach, 5 m (21 12'45"S 159 49'39"W), 2
9, 20.ix.2004; Mangaia: S Ivirua, on Makatea,
30 m (21°55'52"S 157 52'42"W), 1 9, 26.ix.2004;
Ivirua, on Makatea, 30 m (21 55'15"S
157 53'21"W), 4 9, 27.ix.2004.

Megachile (Callomegachile) umbripennis F.
Smith, 1853

Distribution. — Widely distributed in
Southeast Asia and large parts of the South
Pacific (Krombein 1950, Snelling 2003). In
Hawaii it was present as early as mid-19th
century suggesting an introduction by
Polynesians prior to European arrival
(Snelling 2003). Probably introduced to
the Cook Islands by ancient Polynesians
(Pauly and Munzinger 2003).

Previous records are from Rarotonga
(iii.1979, leg. N.L.H. Krauss; 1999, leg.
British Executive Service Overseas), Atiu
(1976, leg. N.L.H. Krauss), Aitutaki (1960,
1977, leg. N.L.H. Krauss; Krauss 1961) and
Pukapuka (ii.2004, leg. G. McCormack)
(McCormack 2004).

Biology. — One female and three males
carry some phoretic mites. This species
nests in all kinds of cavities of suitable size
but with a preference for wood. Plant
resins are used to line the nest cells
(Snelling 2003). This species is perhaps
oligolectic on Fabaceae. The few observed
females frequently visit the native Beach
Pea (Vigna marina (Burnt.)). Males regular-
ly visit the flowers of the introduced
Mexican Fire-Plant (Euphorbia cyathophora
J. A. Murray) (Euphorbiaceae) for nectar.

Specimens examined. — Rarotonga: Arorangi,
upper beach, 5 m (21 14'20"S 159 49'40"W), 15
;, 15.ix.2004; 1 9, 5 $, 18.ix.2004; Arorangi,
fallow land, 10 m (21T2'54"S 159 49'42"W), 1 9,
T-Ux.2004; Arorangi, upper beach, 5 m
(21 12'45"S 159 49'39"W), 2 9, 3 3, 20.ix.2004;
3 6\ 21.ix.2004; Mangaia: Ivirua, on Makatea,
30 m (21 55'15"S 157 53'21"W), 1 3, 22.ix.2004; S

Ivirua, on Makatea, 30m (21 55'52"S
157 52'42"W), 1 9, 1 S, 26.ix.2004.

Megachile (Eutricharaea) sp. 1

Distribution. — From the Australian and
South Pacific region 66 taxa of the sub-
genus Eutricharea are described that can be
divided into three groups (Michener 1965).
This small species belongs to group A as
defined by Michener (1965). It comprises
36 taxa that are badly in need of revision
before the species can be identified. The
male is characterised by a mandible with
an inferior basal projection, front coxa with
apical spine, anterior tarsus slightly broad-
ened and carina of sixth tergum with
median emargination almost obscured by
coarse serration. Eutricharea is widely dis-
tributed in the South Pacific with at least
some species indigenous to its islands. First
record for the Cook Islands of this possibly
indigenous species.

Biology. — The males were patrolling the
upper beach over sparsely vegetated areas
covered with the native Beach Pea (Vigna
marina (Burm.)) (Fabaceae). Most species of
the genus Megachile make nest cells from
pieces cut from leaves of plants (Michener
1965); others line their cells with plant
resins (Michener 2000).

Specimens examined. — Rarotonga: Arorangi,
upper beach, 5 m (21 12'45"S 159 49'39"W), 1
9, 4 $, 20.ix.2004; 3 $, 21.ix.2004; 3 J, 4.X.2004.

Megachile (Eutricharaea) sp. 2

Distribution. — As for the previous spe-
cies, this one belongs to group A of
Michener (1965). This species is unusually
large (female 11-13 mm, male 10-12 mm)
for a member of group A. The male is
characterised by a mandible without an
inferior basal projection, front coxa with
short apical tubercle, anterior tarsus un-
modified and carina of sixth tergum with
large median emargination. For general
comments on the distribution of Eutrichar-

JOURNAL OF HYMENOPTERA RESEARCH

aea see previous species. First record for the
Cook Islands. Perhaps introduced.

Biology. — One male carries a few indi-
viduals of a phoretic mite. For data on
nesting see previous species. This species is
very probably oligolectic on Fabaceae. The
females can be regularly observed to collect
pollen on the native Beach Pea {Vigna
marina (Burm.)) and sometimes on the
introduced Streaked Rattlepod (Crotalaria
pallida Aiton).

Specimens examined. — Rarotonga: Arorangi,
upper beach, 5 m (21 14'20"S 159 49'40"W), 1
9, 15. ix. 2004; Arorangi, upper beach, 5 m
(21T2'45"S 159 49'39"W), 14 9, 3 3, 20.ix.2004;
11 9, 5 3, 21.ix.2004; 5 9, 3 $, 4.X.2004; Mangaia:
Ivirua, on Makatea, 30 m (21 55'15"S
157 53'21"W), 6 9, 2 S, 22.ix.2004; 1 9, 2 J,
27. ix. 2004; Ivirua, Taro gardens, 30 m
(21 55'20"S 157 53'53"W), 1 9, 1 $, 23.ix.2004.

Apidae

Apis mellifera Linnaeus, 1758

Distribution. — Now distributed world-
wide and introduced to the Cook Islands.
It probably occurs on all larger islands and
is known from Rarotonga, Mangaia, Aitu-
taki and Atiu and definitely missing on
Tongareva (= Penrhyn) and Pukapuka
(Krauss 1961, McCormack 2004).

Biology. — The honeybee is highly social
and polylectic. Beside the hives of bee-
keepers it can build nests in hypergaeic
cavities like hollow trees.

Specimens examined. —
Ceratiua (Neoceratina) dentipes Friese, 1914

Distribution. — Japan, southeast Asia,
Australasia and the West Pacific. The Cook
Islands are far outside of the known
distribution of C. dentipes and the subgenus
Neoceratina on the whole (Michener 1965,
Hirashima 1971). The species is recorded
for the first time on the Cook Islands and is
surely introduced.

Biology. — The species is known to build
nests in pithy stems. Nesting biology is

Volume 15, Number 1, 2006

Table 1. Distribution of bees and wasps for each island in the Cook Islands. Data about island tvpe, land area
and maximum elevation after Hein et al. (1997). Codes for islands and abbreviations are as follows: PN:
Penrhyn, RK: Rakahanga, MH: Manihiki, PK: Pukapuka, NS: Nassau, SW: Suwarrow, PL: Palmerston, AK:
Aitutaki, MN: Manuae, MT: Mitiaro, TK: Takutea, AT: Atiu, MK: Mauke, RR: Rarotonga, MG: Mangaia; A: Atoll,
R: Reef Island, M: Makatea, V: High Volcanic Island, n.a.: no data available.

Northern Cook Islam

Southern Cook Island;.

RK MH PK

MT TK AT

Island type

AAA

A/V

M R M

Land area (km2)

9.8

3.9 5.4 3.8

1.1

0.4

1.1

18.0

5.8

30.0 1.4 29.0

18.0

67.0

51.0

Max. elevation (m)

low

low 5 6

low

124

11 6 70

653

169

Species

Chrysis spec.

Anterhynchium rufipes

Delta esuriens okinawae

Pachodynerus nasidens

Parodi/nerus bicinctus

X X

Polistes jokahamae

Polistes olivaceus

Ampulex compressa

Pison spec. 1

Pison spec. 2

Sceliphron caementarium

Sceliphron laetum

Tachysphex f. fanuiensis

Apis mellifera

Cerati)ia dentipes

Litliiirgus scabrosus

Megachile umbripennis

Megachile spec. 1

Megachile spec. 2

Total species no.

n.a.

n.a. n.a. 3

n.a.

1 n.a. 6

analysed in detail by Okazaki (1992). The
bee was found nesting in twigs in a small
forest clearing in the interior of Rarotonga
and on the upper part of the beach where it
was flying above the creepers of the Beach
Pea (Vigna marina (Burm.)) (Fabaceae).

Specimens examined. — Rarotonga: Arorangi,
Raemaru trail, 50 m (21 14'06"S 159 49' WW),
5 9, 17.ix.2004; Arorangi, upper beach, 5 m
(21 12'45"S 159 49'39"W), 2 9, 21.ix.2004.

DISCUSSION

Fauna and biogeograpln/ of the Aculeata of
the Cook Islands. — Information about the
bee and wasp fauna is only available for
seven of the fifteen Cook Islands (Table 1).
No records exist for the islands of Palmer-
ston, Manuae and Takutea in the Southern
Group and Penrhyn, Manihiki, Rakahanga,

Nassau and Suwarrow in the Northern
Group. During my three weeks' stay on
Rarotonga and Mangaia six species were
recorded for the first time (Pison sp. 2,
Sceliplnwi laetum, Ceratina dentipes, Megachile
sp. 1, M. spec. 2) adding up to a total of 19
species of bees and aculeate wasps current-
ly known from the Cook Islands. Except for
the probably extinct Anipnlex compressa and
Chrysis sp. all previously collected species
were recorded again. Among the newly
recorded species are the conspicuous wasp
Sceliphron laetnm and the large and common
bee Megachile sp. 2. This is a clear sign that
further investigations especially on the out-
er islands will probably lead to the discov-
ery of additional species.

From their biology and distribution it is
concluded that probably only three species

34 Journal of Hymenoptera Research

are definitely indigenous to the Cook tigation might show that the real number

Islands: Anterhynchium rufipes, Parodynerus of species on this island is much higher

bicinctus, Tachysphex fanuiensis. All of them than the nine recorded species and that it

are widely distributed in the tropical would fit into this scheme as well. A lack of

Pacific. Three more species (Pison sp. 1, correlation between species diversity and

Pison sp. 2, Megachile sp. 1) might be size of an island could also be predicted

indigenous or even endemic because many from the low number of (probably) in-

of their relatives are known to be restricted digenous species that indicates an under-

to other Pacific islands (Yasumatsu 1953, saturation of its fauna, probably due to

Michener 1965). But their status remains their isolation.

uncertain due to the lack of revisions. Of Biogeography of the aculeate fauna of
the remaining 13 species 11 were accident- Pacific islands. — The biogeography of the
ly introduced. The origin of eight species is bee and wasp fauna of the Cook Islands
in southeast Asia, two were introduced can only be understood in a Polynesian
from the Americas and the provenance of and Pacific context. Traditionally the Pa-
Chrysis spec, is unknown. All these species cific islands, excluding those close to the
build their nests overground in pre-exist- Americas, Asia and Australia, are divided,
ing cavities, dead wood and pithy stems, originally on the basis of their indigenous
construct free clay nests or free hanging aboriginal peoples, into Polynesia, Micro-
paper nests. The only exception is the nesia and Melanesia (Crocombe 2001). This
cuckoo wasp Chrysis sp. that very probably division serves sufficiently for biogeo-
parasitises a wasp species with hypergaeic graphic purposes (Fosberg 1984) and is
nests. Thus, the nests are easily transported used here, too. Good accounts of the bees
and species have been carried to various and wasps are available for very few of the
islands (Michener 1965). Of the six possibly Pacific islands or island groups (Table 2).
indigenous species five build overground Therefore, biogeographic conclusions on
nests, too, and only T. fanuiensis is a ground their island faunas must be regarded as
nester. In addition to the honeybee only the tentative except at the broadest level,
digger wasp A. compressa was intentionally From Polynesia faunistic data on bees
introduced to control the cockroach Peri- and wasps are available for seven archi-
planeta americana (Walker and Deitz 1979). pelagos including New Zealand. The most
With increasing tourism and trade acci- comprehensive investigations were made
dental introductions of additional species on six islands groups of Micronesia whilst
can be expected in the future. Melanesia is poorly investigated (Table 2).
For the comparatively well investigated As demonstrated for Micronesia (Krom-
islands Pukapuka, Aitutaki, Mangaia and bein 1950) there is a general tendency that
Rarotonga a positive relationship exists the number of species as well as endemics
between island size and number of species of Pacific islands declines with increasing
(Table 1) as predicted by the theory of distance from Australian and Asian main-
island biogeography (MacArthur and Wil- lands and the Indo-Australian islands,
son 1967). But this relationship is probably Thus, most of the remote archipelagos of
artificial and caused by accidental intro- Polynesia possess a depauperate aculeate
ductions. Except for the very touristic fauna consisting of widespread species.
Aitutaki the other three islands show This, too, applies to the indigenous acule-
a correlation of species diversity and in- ate fauna of the Cook Islands that is
tensity of traffic that facilitates introduc- extremely depauperate and only consists
tions. While the intensity of faunistic of three wasp species widely distributed in
surveys on Aitutaki is low compared with Polynesia and other parts of Oceania. Due
the aforementioned islands further inves- to a lack of taxonomic revisions the

Volume 15, Number 1, 2006

Table 2. Numbers of (sub)species of aculeate wasps and bees (including introduced species) in Polynesia
and adjacent islands of Melanesia and Micronesia (n.a.: no data available; (): species numbers for some families
not available).

Apidae

"Sphecidae'

Vespidae

Pompilidae

Scoliidae

Mutillidae

Tiphiidae

Chrvsididae

total

endemic

introd.

Polynesia

Hawaii I)2)3)

127

275

76%

24%

Samoa 4)5|6)

n.a.

Marquesas Is.7>22)23)

n.a.

Tuamotu Is.22)23)

n.a.

Society Is.22'23*25'

n.a.

Cook Is.

84%

New Zealand 8,27)28)

>40

n.a.

(74) (

±80%)

n.a.

Melanesia

Solomon Is. 9,10)

n.a.

(32)

n.a.

Vanuatu ,1,12)13)

n.a.

(14)

n.a.

New Caledonia

n.a.

14)21)24)26)

Fiji 15)16)

n.a.

Micronesia

Bonin Is. 17)291

47%

Northern Marianas

25%

42%

18)19)20)

Southern Marianas

39%

42%

18)19)20)

Carolines 18,,9)2n>

50%

25%

Marshall Is. ,8)19)20>

55%

Gilbert Is. ,8)

100%

"Snelling (2003), 2lDaly & Magnacca (2003), 3,Nishida (1994), 4lPauly & Munzinger (2003), S| Perkins &
Cheesman (1928), h) Williams (1928), 7> Williams (1932), 8) Donovan (1983), 9) Krombein (1949a), ""Krombein
(1951), '"Cheesman (1936), 12)Cheesman & Perkins (1939), ,3)Cheesman (1948), 141 Pauly & Munzinger (2003),
l5) Williams (1947), 16,Fullaway (1957), 17)Yasumatsu (1955), m Krombein (1949b), 19) Krombein (1950), 20,Tadauchi
(1994), 211Callan (1990), 22,Michener (1965), 23) Cheesman (1928), 24> Turner (1919), 25)Menke (1979), >l Williams
(1945), 271 Harris (1987), M> Harris (1994), 29)Yasumatsu (1936)

recorded species of Pison and Megachile
(Eutricharaea) are unidentifiable at present
but they are probably not endemics of the
Cook Islands. Exceptions are Hawaii with
its highly endemic and species rich fauna
that result from the adaptive radiation of
genera like Hylaeus (Nesoprosopis) (60 spe-
cies) and Odynerus (99 species) and the
incompletely documented fauna of New
Zealand. There is also a rapid eastwards
loss of indigenous parasitic taxa of the
families Scoliidae, Mutillidae, Tiphiidae
and Chrysididae. They are all dependent
on their hosts and thus have a limited
potential for dispersal. Interestingly, this
also seems to be true for the spider hunting
Pompilidae.

These basic patterns are nowadays at
least partly obscured by the number of

introduced species that often comprise up
to 50% or more of the species known to
occur on some archipelagos (Table 2). As
a basis for the reconstruction of distribu-
tion patterns and to better understand the
history of island colonization in the Pacific
there is a need for further investigations of
island faunas and taxonomic revisions of at
least the most diverse and widely distrib-
uted genera of bees and sphecid wasps.

ACKNOWLEDGEMENTS

I am very much indebted to Gerald McCormack,
Director of the Cook Islands Natural Heritage Project,
for his constant support and for providing valuable
background information about the bees and wasps
recorded in the Cook Islands Biodiversity Database.
Species identifications were carried out by (in alpha-
betical order) James Carpenter (New York), Michael

Journal of Hymenoptera Research

Ohl (Berlin), Wojciech Pulawski (San Francisco) and
Michael Terzo (Gembloux), who also provided in-
formation about nomenclature and distribution. Field
work was very much facilitated by the help of Ulrike
Gigengack (Minister). The National Research Com-
mittee, Government of the Cook Islands, kindly
permitted the collection of specimens on Rarotonga
and Mangaia. A previous version of the manuscript
was greatly improved by the comments of Wojciech
Pulawski.

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J. HYM. RES.
Vol. 15(1), 2006, pp. 38-163

A Review of the New World Genera of
Trichogrammatidae (Hymenoptera)

John D. Pinto

Department of Entomology, University of California, Riverside, Riverside, CA 92521, USA

john.pinto@ucr.edu

Abstract. — The 55 New World genera of Trichogrammatidae are reviewed and a key is provided
for their identification. The family classification utilized recognizes two subfamilies and four tribes:
Trichogrammatinae (Trichogrammatini) and Oligositinae (Paracentrobiini, Chaetostrichini, and
Oligositini). Four new genera are proposed, Adelogramma, Pseuduscaua, Thanatogramma, and
Viggianiella. Five generic group names are given renewed status. These include Burksiella De
Santis, Centrobiopsis Girault, Ceratogramma De Santis, and Zaga Girault, at the genus level, and
Lutzimicron Costa Lima, at the subgenus level. Four genera are placed in synonymy: Uscanoidea
Girault (= Gnorimogramma De Santis), Zaga (= Lathrogramma De Santis), Burksiella ( =
Parahispidophila Yousuf and Shafee), and Xenufens Girault (= Pseudoxenufens Yoshimoto).
Fifteen new or revised species combinations result from generic synonymy and resurrection. Nine
new species described include Adelogramma pritnum, Burksiella dianae, Ceratogramma jeffersi,
Lathromeris hesperus, Lathromeroidea exemplum, Lathromeroidea gerriphaga, Pseuduscaua sola,
Thanatogramma oweni, and Viggianiella tropica.

The chalcidoid family Trichogram-
matidae represents a ubiquitous group of
insect-egg parasitoids that is among the
most poorly known of Hymenoptera. Min-
ute size, fragility and logistical problems
associated with collecting and curating
such insects are largely responsible for
their uncharted taxonomy and biology
(Pinto and Stouthamer 1994). Yet, since
the benchmark family review by Doutt and
Viggiani (1968) there has been advance in
our understanding of the group. A large
number of new genera have been added,
generic concepts have been modified and
there has been a significant increase in
collections available for study. This work is
an attempt to consolidate current knowl-
edge associated with the New World
genera of Trichogrammatidae. Included
are a key and review of genera, and
descriptions of new genera and species.

Current address: P.O. Box 2266, Waldport, OR 97394,
USA; (e-mail: john.pinto@iRr.cdu)

In addition, certain nominal genera are
synonymized and others are returned to
generic status.

Although poorly known everywhere,
knowledge of New World trichogramma-
tids has lagged and continues to lag. The
last comprehensive review of genera
(Doutt and Viggiani 1968) reported only
28 in the New World, or about 40% of the
total known at that time. This study
recognizes 55 New World genera, over
60% of the current total of 88 extant genera.
The geographic distribution of these gen-
era is summarized in Table 1.

It is the generic level, where modest
advances have been made, that is the focus
of this study. Species taxonomy remains
especially rudimentary. Of the 232 named
New World species, over 40% belong to
a single genus, Trichogramma, a group
closely tied to biological control (Pinto
1999). Several other relatively large genera
such as Oligosita, Pseudoligosita, Mirufens,
and Chaetostricha have few or no species

Volume 15, Number 1, 2006 39

named in the New World, although they only from the Western Hemisphere will be
are relatively diverse there. Other groups, shown to have broader distributions once
such as Burksiella, Zagella, Lathromeroidea, the family is more adequately sampled.
Uscanaidea and Zaga in particular, are Considering that large areas of the New
especially speciose in the Western Hemi- World, especially in South America, re-
sphere, yet remain almost completely un- main poorly collected, the key to genus
known. will be considered successful if 80-90% of

Historically, the greatest impediment to the collections can be successfully identi-

progress in trichogrammatid taxonomy has fied.
been the absence of adequate collections.

Until recently, holdings of most genera MhlHODS
have been restricted to types and, at best, Collecting and curation. — Trichogram-
a few badly curated specimens other than matidae are collected by a variety of
types. In many cases, the types themselves methods. Unfortunately relatively few are
are in such poor condition that the taxa taken directly from host, although Tricho-
represented are unidentifiable. Most of the gramma and Trichogrammatoidea are excep-
recent advances can be attributed to the tions because of association with biological
marked increase in survey activity by control projects and frequent occurrence on
parasitic hymenopterists in the last three hosts (Lepidoptera) which are easily de-
decades. The University of California Riv- tected and abundant in disturbed habitats,
erside Collection has served as a primary Most material, however, is removed from
depository for uncurated Tricho- bulk arthropod samples taken in yellow
grammatidae resulting from much of this pan traps (YPT), fine-meshed Malaise traps
work (see Acknowledgments). Although (MT), flight intercept traps (FIT), or sweep
material is received from all parts of the samples (SP). Noyes (1982) reviewed these
world, collections from both temperate and methods for chalcidoid collecting. For this
tropical areas of the Western Hemisphere study most specimens collected were
have dominated. Except for types, limited stored in 80% ethanol and placed in
historical material in various museums and a refrigerator or freezer until they could
certain literature records, this study is be curated further.

based almost totally on these recent accu- Taxonomic studies of Trichogram-

mulations. matidae require slide-mounted material

Any attempt to review the trichogram- and examination with the light microscope

matid genera for an extensive region of the at magnifications up to at least 600X.

world deals with a moving target. The Specimens utilized here were mounted in

family is a long way from attaining Canada balsam following the techniques of

taxonomic stasis and new genera are Platner et al. (1998). Most were placed

expected as collecting continues. Also, dorsoventrally on slides. For certain taxa

because the vast majority of species in the lateral mounts were used as well. Slides

family remain undescribed, generic defini- were complemented by card-mounted in-

tions and limits will be prone to revision dividuals whenever possible to allow

for some time. It is hoped that reviewing examination of color, body length and

the New World genera now will encourage shape. Specimens to be carded were re-

and facilitate the work that remains, moved from ethanol and treated with

Although this review focuses on the New hexamethyldisilazane (HMDS) to prevent

World for practical reasons, it should be shrivelling (Heraty and Hawks 1998).

emphasized that most of its genera also are Scanning electron microscopy was used

extralimital in distribution (Table 1). In- frequently to fully appreciate structure. For

deed, it is likely that certain genera known SEM work, specimens were taken from

Journal of Hymenoptera Research

Table 1. General geographic distribution of New World genera of Trichogrammatidae.

Genus1

North America

Central America

West Indies

South America

Extralimital

Adelogramma

Adryas

Aphelinoidea

Bloodiella

Brachista

Brachygrammatella

Brachyufens

Burksiella

Centrobiopsis

Ceratogramma

Chaetogramma

Chaetostricha

Doirania

Epoligosita

Haeckeliania

Hydrophylita

Ittys

Ittysella

Lathromeris

Lathromeroidea

Megaphragma

Mirufens

Monorthochaeta

Nicolavespa

Oligosita

Pachamama

Paracentrobia

Paratrichogramma

Pin ton

Poropoea

Prestwichia

Pseudoligosita

Pseuduscana

Pterandrophysalis

Pteranomalogramma

Pterygogramma

Sinepalpigramma

Soikiella

Thanatogramma

Trichogramma

Trichogrammatella

Trichogrammatoidea

Trichogrammatomyia

X
7

Tumidiclava

Tumidifemur

Ufens '

Uscana

LI scan ell a

Uscanoidea

Usomopsis

Viggianiella

X
X

Xenufens
Xiphogramma

X
X
X

Zaga

Zagella

Volume 15, Number 1, 2006

ethanol or off of cards and similarly treated
with HMDS (Nation 1983).

Types. — The deposition of primary types
and paratypes of new species is indicated
in the descriptions. To minimize the
possibility of mixed type series paratype
designations generally are restricted to
topotypic material. Mixed series are a con-
cern in the Trichogrammatidae where in-
terspecific morphological differences are
often minimal and may be lacking alto-
gether (Pinto et al. 2003).

Types of type species were examined for
the majority of nominal genera treated.
Specifics are indicated in the text when
necessary to justify proposed taxonomic
changes.

Generic key. — The generic key is intended
for identification of New World taxa only.
Although both sexes are considered in the
key, females will produce better results in
most cases. Ideally both sexes will be
available for identification. Females are
known for all described genera. Males are
known for all except Pteranomalogramma,
Sinepalpigramma, Uscanella and Viggianiella.
Because of strong sexual dimorphism,
considerable intrageneric variation or an-
ticipated character misinterpretation cer-
tain genera terminate in the key more than
once. The key assumes the availability of
cleared, slide-mounted specimens, and use
of a light microscope with adequate mag-
nifications.

Generic reviews. — A synopsis of each
genus is provided. Included are a synony-
my with all relevant names, a diagnosis,
a summary of general distribution, com-
ments on worldwide and New World
diversity, a discussion of nomenclature
and taxonomy, New World records, and
host information. For most genera the
diagnosis is based on females. For males,

only genitalic features are reported in
many cases. It can be assumed that males
do not differ from females for other
characters unless indicated. Characters
stressed in each diagnosis focus on those
important for distinguishing the genus
from similar taxa, and consequently are
not homogeneous. References for original
descriptions are not included in the Liter-
ature Cited unless cited in another context.
This primary literature is referenced by
Doutt and Viggiani (1968) and Noyes
(2001, 2005).

The records for each genus include
country and, for countries occupying con-
siderable latitude and /or longitude, state,
province or region as well. These addition-
al categories are reported for Canada, The
United States, Mexico, Brazil, Argentina
and Chile. There is no attempt to provide
detailed host information. Several pub-
lished host records of trichogrammatids
lack adequate documentation and are
suspect. In this work host family or order
is provided (with at least one reference)
only if the record is considered trustwor-
thy.

Generic index. — A formal index to genus
is not provided. However generic treat-
ments can be found in the text by assigned
numbers which are based on order of
presentation. These are given in the classi-
fication section below, and in the generic
key.

Species descriptions. — Descriptions of new
species are kept to a minimum in this
work. Only nine are described. Included
are type species of the four new genera,
and five species which represent relatively
distinctive sections of their genus. The
species descriptions are included in the
treatment of the appropriate genus. Sec-
tions for each new species treatment are

1 The record of the Australasian genus Bracln/ia in the New World by De Santis (1997) is in error. See treatment of
Zagella [48].

42 Journal of Hymenoptera Research

the description, types, etymology, distri- Trichogrammatid structure has never
bution, material examined and comments, been studied in detail and it largely
The material examined section also in- remains an untapped resource to be
eludes an abbreviated listing of the types, exploited for infrafamilial systematics.
The comments section includes compari- One problem precluding facile utilization
sons to related species when appropriate, of structure obviously is the small size
Quantitative data are taken from the type (usually < 1.0 mm) and fragility of these
series in most cases. wasps. Slide-mounted material, so neces-
Museums— Acronyms for museums re- sary to characterize antennae, wings and
ferred to in the text follow N. Evenhuis and genitalia, generally precludes detailed
G. Samuelson (Abbreviations for Insect study of core body parts. For this reason,
and Spider Collections of the World: slide mounts need to be complemented
http://hbs.bishopmuseum.org/codens/ with SEM examination before morpholog-
codens-inst.html). These are as follows: ical variation in this family is adequately
BMNH (Natural History Museum; Lon- understood. The following is a summary of
don, UK), CAS (California Academy of the structures (with acronyms) character-
Sciences), CNC (Canadian National Collec- izing the family and used for identifying
tion of Insects, Ottawa), DEZA (Diparti- and diagnosing genera,
mento di Entomologia e Zoologia Agraria Head capsule (Figs 163-166). — The head
dell'Universita, Portici), EMEC (Essig Mu- capsule is used minimally in this study,
seum of Entomology, U.C. Berkeley), HIC The extent of sclerotization of the head
(Hymenoptera Institute Collection, Univer- varies. In general, the head is poorly
sity of Kentucky), INBC (Instituto Nacional sclerotized except for the gena. However,
de Biodiversidad [INBio], Costa Rica), a sclerotized plate may occur on the vertex
INHS (Illinois Natural History Survey, and in a few groups the face also is well
Champaign), MACN (Museo Argentino sclerotized. Subantennal grooves (SAG)
de Ciencias Naturales "Bernardino Riva- are characteristic of most or all genera,
davia"), MLPA (Museo de la Plata, Argen- whereas the presence of malar sulci (MSL)
tina), QM (Queensland Museum, Austra- varies. A distinct clypeus (CLY) usually is
lia), MZCR (Museo de Zoologia, Universi- present but there are exceptions. The
dad de Costa Rica), NMNH (National placement of the toruli (TOR) on the face
Museum of Natural History, Washington varies somewhat and can be useful in
D.C.), TAMU (Texas A&M University, distinguishing certain genera.
College Station), UCDC (University of Antenna (Fig. 1).— The antenna is of
California, Davis), UCRC (University of considerable importance in trichogramma-
California, Riverside). tid systematics. Of primary use is the

number and arrangement of flagellar see-

ANATOMICAL STRUCTURE ments (thoS(? . * ^ & [s] Jd

AND TERMINOLOGY pedice| [p]) Xh can vary from two (in

Anatomical terms for most general Hy- male Trichogramma) to nine (in certain male

menoptera structures follow Gibson (1997). Ceratogramma). The first segments (proper-

Doutt and Viggiani (1968) is utilized for ly subsegments) beyond the pedicel are the

terms associated with the fore wing. Those ring-like anelli (A). There commonly are

used for male genitalia follow Viggiani two anelli (Al, A2), although several

(1971) and Pinto (1999), and those applied genera have only one, and in Ceratogramma

to antennal sensilla are consistent with there are three (A3). In many cases A2 is

Pinto and Owen (2004). References to partially or almost completely fused to the

length and width of structures refer to following segment. The presence of a coe-

maximum dimensions unless indicated. loconic sensillum on A2 (visible medially)

Volume 15, Number 1, 2006

■

FrS

. • . . .

Figs 1-4. 1, antenna, Ceratogramma brasiliense (9). 2, fore wing, Haeckelmnia. 3, hind wing Ceratogramma masneri
(arrow to hamuli). 4, same, Ufens. See text (Anatomical Structure and Terminology) for explanation of acronyms.

identifies this segment (Fig. 10). Flagello-
meres apical to the anelli are referred to as
postanellar segments. They either are con-
solidated into a club (C) of one to five
segments (C1-C5), or also partitioned into
a funicle of one or two segments (Fl, F2).
When a funicle is present the club gener-

ally is one to three segmented, although
a small fourth segment occurs in males of
Ufens, Mirufens and some Ceratogramma
(Fig. 60). In some cases deciding whether
a segment represents a funicular or a club
segment is difficult and somewhat arbi-
trary. Also, in certain taxa two or more of

44 Journal of Hymenoptera Research

the flagellomeres can be partially fused characterized after SEM study by Olson

resulting in segment counts that vary with and Andow (1993). Their identification in

the surface viewed. The sutures separating most other genera is based simply on

flagellar segments also vary considerably phenetic similarity to those in Tricho-

in form. They may be transverse but, more gramma.

commonly, are slightly to highly irregular Regarding the distribution of sensilla on

resulting in noticeable segment asymme- the flagellar segments: PLS, APB, APA,

try. This asymmetry results in segment BPS, and FS can be found on any of the

length varying with the surface being postanellar segments depending on the

viewed. group. RS occur apicoventrally on the last

Because of their considerable size range, club segment of females in certain genera,

frequent partial or complete fusion and They may be numerous and comprise

considerable asymmetry, segment number a distinct patch or 'micropilose region' as

is often difficult to determine. Small seg- in Trichogramma (Fig. 34), or be represented

ment size is especially problematic. Thus, by no more than two-three sensilla near the

in several genera such as Zagella, Adryas, apex of the club as in certain Adryas. RS are

and Zaga Fl or CI is narrow, very short, similar to FS except they are shorter,

transverse, and closely appressed to the recurved and have characteristic oblique

following segment (F2 or C2). In these taxa, surface grooves (Pinto and Owen 2004,

this segment is difficult to detect or may be Fig. 8). These grooves are not visible with

confused with an anellus. Fortunately, in the light microscope; consequently RS in

most cases Fl or CI will bear one or more most cases are tentatively identified by

basiconic peg sensilla on its apical margin their length, position and curvature. A

(Figs 5, 6); anelli never do. It often is the single CS occurs on A2 (Fig. 10) and one

detection of these sensilla that signals the sometimes can be detected on F2. UPP

presence of a basal Fl or CI. Locating these occur singly at the apex of the club in

segments is critical for generic identifica- females (Fig. 11); it is not clear if they are

tion. Although segments may also be present in all genera.

partially fused, in this work segment In females of several genera the apex of

number refers to completely divided seg- the club is abruptly narrowed and pro-

ments unless indicated. longed into a terminal process (= apical

Because of their value in identification process of Doutt and Viggiani 1968, and
the distribution of antennal sensilla are Pinto and Owen 2004) of varying length
reported in many of the generic diagnoses (TP, Fig. 11). This structure is not to be
and species descriptions. Exemplars of the confused with PLS which sometime exten-
various sensilla associated with tricho- sively overlap the club apex (e.g. Fig. 89).
grammatid antenna are given in Figs 5- When present, the terminal process usually
11. Included are placoid sensilla (PLS), extends considerably beyond the tip of the
aporous sensillar trichodea B (APB), apor- apical-most PLS and, as is typical of the
ous setae A (APA), flagelliform setae (FS), female club in most genera without a pro-
recurved sensilla (RS), basiconic peg sen- cess, bears the UPP at its apex,
silla (BPS), uniporous pit pore sensilla In addition to presence or absence and
trichodea D (UPP), and coeloconic sensilla density on various segments, sensilla
(CS). These terms and acronyms (also shape and size also are useful taxonomi-
utilized in Pinto and Owen 2004) follow, cally. The various setiform sensilla may
or are modifications of, those utilized for vary considerably in length, and the BPS
Trichogramma by Vincent and Goodpasture vary in shape from subglobose to narrow
(1986), Olson and Andow (1993), and Pinto and fusiform. Also, the degree of attach-
(1999). Most of these structures were ment of the longitudinal PLS to the

Volume 15, Number 1, 2006

Figs 5-11. Antennal sensilla. 5, Zagella. 6, Lathromeroidca. 7, Soikiella occidentalis ( J). 8, Soikiella occidentalis (9). 9,
Trichogramma pretiosum (9). 10, Tumidiclava. 11, Nicolavespa theresae (9). See text (Anatomical Structure and
Terminology) for explanation of acronyms.

46 Journal of Hymenoptera Research

antennal surface varies. In most taxa they family but have yet to be studied ade-

are attached almost their entire length with quately.

only the apex projecting beyond the sur- Mesosoma. — Except for the wings, rela-

face (Fig. 6). However, these sensilla may tively few characters of the mesosoma are

be free of the surface over much of their used for identification. Among these

length, and in males of certain genera (Figs 179-194) are the number of setae on

(Soikiella, Haeckeliania, Trichogrammatoidea, the midlobe of the mesoscutum (MLM)

Adelogramma), they are completely free, and scutellum (SCT), shape and length of

attached only at their base, and project the propodeum (PPD), relative length of

from the antenna as thick, elongate spini- the metanotum (MTM), and presence or

form structures (Fig. 7). Consequently they absence of the mesopleural suture (PS) and

may be confused with FS, the most transepisternal sulcus,

common setiform sensilla on the antenna. There are one or two pair of setae on

Unlike the relatively smooth and thin both the midlobe and scutellum of most

FS, however, these exserted PLS are genera (Figs 180, 184-187). All representa-

thicker and retain their characteristi- tives of the Oligositini have a single pair on

cally ridged and densely pitted surface both structures. In all other groups at least

(cf. Figs 6-8). the scutellum has two pair, although the

Mouthparts. — Relatively few features of anterior pair may be very small and

the trichogrammatid mouthparts have difficult to see (Figs 184, 185). In a few

been used in taxonomy. Among these are cases (e.g. Brachyufens, Fig. 179) there are

the number and structure of the mandib- three pair of midlobe setae, and in one

ular teeth and the number of maxillary species of Pierygogramma, an anomaly in its

palpal segments. Mandibular dentition is genus as well as the family, the scutellum

difficult to determine on slide-mounted has six pair (Fig. 95). The number of setae

specimens. Most genera have a mandible on the midlobe generally is constant within

with three-five serrations or teeth (Fig. 168). genera, however exceptions occur (e.g. in

The mandibles are somewhat palmate in Adryas, see Pinto and Owen 2004). The

structure with the teeth arranged antero- shape of the propodeum and its medial

posteriorly when the head is in a hypo- length relative to that of the metanotum

gnathous position. In a small group of often is reported. The central section of the

genera which includes Trichogramma there propodeum may be undifferentiated or

is a distinct anterior socketed tooth as well elevated and expanded posteriorly (propo-

(Fig. 169). This appears to be analogous to deal disk, Figs 186, 187).

the ventroapical socketed mandibular The presence of sutures and sulci on the

tooth found in Aphelinidae and Encyrtidae mesosoma usually are difficult to detect in

(Heraty and Schauff 1998). There typically trichogrammatids without SEM examina-

are one or two maxillary palpal segments tion. This is not only because of small body

in trichogrammatids (Figs 172, 173). A peg- size but also because the relatively thin

like or setiform sensillum accompanied by cuticle often reveals subsurface ridges that

one-two setae usually occur at or near the are easily confused with surface impres-

apex of the terminal palpal segment, sions. For example, what appears to be

Although the palp may be quite small, a pleural suture under the light microscope

only in one genus, Sinepalpigramma, is it may only be the internal pleural ridge,

entirely absent (Fig. 171). The palpal Similarly, what appear on slides to be

segment count may be critical for generic distinct axillae (AX) and axillulae (AXL)

identification. A single segmented labial generally are not actually separated from

palp also occurs in Trichogramma tidae. Its the scutellum as distinct sclerites on the

size and other labial features vary in the surface (Figs 190, 191). These and other

Volume 15, Number 1, 2006

mesosomal structures have yet to be
adequately evaluated for use in family
taxonomy.

Except for occasional reference to tar-
somere length, tibial /femoral spur struc-
ture, and overall shape, leg structure is
used minimally. One important character
however is the presence or absence of
thorn-like spines on the dorsal surface of
the foretibia. A spinose surface charac-
terizes MirufetiS; Chaetostricha and Zaga and
helps separate them from similar genera
(cf. Figs 175-178). A potential leg character
which has not been adequately studied is
the spatulate structures at the apex of the
hind tibia (Fig. 174) and on tarsomeres I — III
of the fore leg. A feature common to
Chalcidoidea, these apparently modified
setae were previously noted on the hind
tibia of Trichogramma (Hung 1990), but they
occur on the fore and hind legs of all
trichogrammatid genera examined with
SEM (Pinto, unpubl.). It is assumed that
they are used in grooming.

Mesosoma: wings. — Fore wing shape, ve-
nation and discal setation vary considerably
in the family. Figs 2 and 148 indicate those
features utilized in the key and diagnoses.
Wing length (FWL) is taken from the apex
of the humeral plate to the wing apex; wing
width (FWW) is the maximum width
wherever it occurs. Both measurements
exclude the fringe setae. The length of the
longest fringe setae (FrS) is given as
a fraction of FWW. The longest setae
generally occur on the posteroapical mar-
gin. Reference is sometimes made to the
retinaculum (Re), that section of the poste-
rior wing margin which the hamuli of the
hind wing (Figs 3, 4) hook on to.

Terms associated with fore wing vena-
tion include the subcostal (SC) (= sub-
marginal), premarginal (PM) (= para-
stigma), marginal (MV) and stigmal (SV)
veins. A small postmarginal vein is rarely
present in trichogrammatids. The SV usu-
ally includes an apical enlargement or
stigma and a constriction between the
stigma and the MV. In several taxa the SV

is highly reduced and sessile to the MV or
homogeneous its entire length (i.e. without
a basal constriction). All veins are entirely
confluent or the SC and PM may be
disjunct from one another and the MV.
The SV always is confluent with the MV.
The PM has one or two campaniform
sensilla at its apex (Fig. 158). PM length is
taken from its base to the apical-most
campaniform except in the few cases
where this sensillum clearly lies basal to
the vein's apex. The length of the MV is
taken from base to apex. If it is confluent
with the PM, its basal end is taken to be
immediately beyond the apical-most cam-
paniform sensillum. The length of the SV is
determined by measuring the entire dis-
tance from the anterior border of the MV to
the apex of the SV and then subtracting the
apical width of the MV. In most cases the SV
emanates abruptly from the posteroapical
corner of the MV and the two veins are
distinct (Fig. 2); in others such as Tricho-
gramma the entire MV gradually curves
away from the anterior wing margin to form
the SV (Fig. 113). The latter is termed
apically sinuate venation. Similarly, the
MV may end abruptly at its base, or curve
dramatically to meet the PM as in Pacha-
mama (Fig. 103) (basally sinuate venation).
The space between the wing margin and the
PM and SC is the costal cell (CC).

A ratio which is used to help separate
certain genera of Chaetostrichini is the
distance from the large globose sensilla
on the SV to the base of the MV divided by
the length of the PM (SV + MV / PM,
Fig. 161). The placement of these sensilla
vary little if at all among genera and
represent the apical-most reach of the
venation in the wing membrane. This ratio
reflects differences in the length of the MV
vs PM but also in the angle between the SV
and MV. A cuticular spur or radial process
(RP) radiating from the base of the PM into
the wing membrane (Fig. 2) is characteris-
tic of several genera.

Setal density and arrangement on the
fore wing disk are described for most

48 Journal of Hymenoptera Research

genera. The family often is characterized a few groups (Figs 262-264). These consist

by the arrangement of discal setae into of patches of modified cuticle or subcuti-

linear tracks (Fig. 2). Scattered setae usual- cular tubiform structures. They are easily

ly occur between the tracks and if these are seen in slide-mounted specimens although

sufficiently dense the tracks themselves are their structure has yet to be studied,

difficult to discern (Fig. 102). In several Metasomal sternal features restricted to

genera, all discal setae are scattered and no males of Eteroligositina (Oligositini) were

linear arrangement is discernible (Fig. 105), recently illustrated and discussed by Pinto

or the linear tracks are restricted to and Viggiani (2004). The extent to which

a portion of the wing membrane. Several the apical sternum, or hypopygium, sub-

of the linear tracks have been named based tends the female ovipositor varies consid-

on their position (Fig. 148). The most erably in the family (Figs 195-197). Al-

important for generic identification is the though hypopygial length usually can be

RSI, a track running posteriorly from the determined on slide-mounted specimens,

stigma. In several genera a distinct RSI is shape variation, also considerable, is more

absent. Appropriate figures should be difficult to document,

consulted for interpreting terms relating Metasoma: ovipositor. — Ovipositor struc-

to fore wing setal density in the descrip- ture varies. Its length (OL) is reported in

tions and diagnoses. In general, Fig. 2 relation to the hind tibial length (HTL).

represents 'moderately densely setose', Other variation, although considerable, has

Figs. 102 and 105 illustrate 'densely setose', not been adequately studied. It appears to

and Fig. 153 shows 'sparsely setose'. be of taxonomic value primarily at the

The hind wing, although structurally intrageneric level,

simple, also may be useful for generic Metasoma: male genitalia. — Male genitalia

identification and diagnosis. Width, ar- are of importance for both identification

rangement of discal setae, and fringe and classification of Trichogrammatidae

length vary. Discal setae may be present (Viggiani 1971, 1984; Pinto 1997a, 1999).

or absent, and if present either scattered or Terms are illustrated in Figs 198, 199.

arranged in one to several complete or Trichogramma is used to demonstrate ter-

partial linear tracks (Figs 3, 4). The length minology because it possesses virtually all

of these tracks, when reported, is taken structures possible. In Trichogramma, as in

from the hamuli to their apex. several genera, the aedeagus (ADG) is

Metasoma: non-genitalic features. — Except distinct from the phallobase or genital

for genitalia, metasomal characters have capsule (GC) and is capable of indepen-

been all but ignored in the trichogramma- dent movement. A pair of apodemes

tid literature, and relatively few are re- (AAP) extends anteriorly from the base of

ferred to here. In several genera the first the aedeagus. Aedeagal length (AL), when

metasomal (petiolar) tergum bears a trans- reported relative to HTL, includes the

verse row of erect denticles on its posterior apodemes; in several cases reference is

margin. This feature, easily seen with the made to the length of the genital capsule

SEM (Fig. 188), also is visible on slide- (GL). Independent movement of the aede-

mounted specimens (Fig. 189). The meta- gus in Trichogramma and other trichogram-

somal terga normally are homogeneously matines is assumed because its position

sclerotized their entire length. In the relative to the genital capsule often varies

oligositine subtribe Eteroligositina, howev- in conspecifics, and the aedeagus is con-

er, the posterior half of certain terga are siderably narrower than the narrowest

longitudinally striate (Pinto and Viggiani width of the capsule and distinctly sepa-

2004) (Figs 187, 197). Modifications of terga rated from its lateral walls. This condition

VI and VII, restricted to males, occur in is referred to here as Type 1 genitalia

Volume 15, Number 1, 2006

(Figs 198-223). In what presumably is the
most generalized state (e.g. Mirufens and
Haeckeliania; Figs 205, 209), the aedeagus
lies within a trough-like capsule which,
perhaps except for a membranous sheath,
is completely open dorsally. In other
genera (e.g. Paratrichogramma, Tricho-
gramma; Figs 198, 210) this dorsal trough
is partially closed above the aedeagus near
the middle of the capsule delimiting a basal
anterodorsal aperture (ADA). Closure is
effected by a transverse band (Fig. 210)
which may be prolonged posteriorly into
a dorsal lamina (DLA) (Fig. 198). Ventroa-
pically the genital capsule may be differ-
entiated into lateral parameres (PAR),
a medial intervolsellar process (IVP) and,
between this process and the parameres,
a pair of volsellae (VS). Digital spines
commonly occur at the apex of the volsel-
lae, and a ventral ridge (VR) may extend
anteriorly from the IVP base or from that
location if an IVP is absent. Parameres
sometimes bear an apical spine, and in
certain genera such as Haeckeliania and
Paratrichogramma they are bifid apically.

In the majority of trichogrammatid gen-
era, the aedeagus appears to be fused to the
genital capsule to a varying degree and no
longer is capable of independent move-
ment. In these genera the position of the
aedeagal apodemes, when present, does
not vary relative to the capsule among
conspecifics and the aedeagus is not clearly
distinguished from the capsule at least not
at its apex. This represents Type 2 genita-
lia (Figs 224-259). As indicated, the apo-
demes of the aedeagus often remain in this
type of genitalia (e.g. Figs 224-227). Para-
meres and /or volsellae do (Fig. 233) or do
not occur (Fig. 243). In the most extreme
consolidation, the aedeagus and genital
capsule are fused into a simple tubiform
structure with no indication of additional
genitalic parts (e.g. Figs 242-245). In this
case, at most, a pair of setae occur ventrally
(ventral setae) (e.g. Figs 247, 253).

In most genera with Type 2 genitalia,
a relatively large aperture (ADA) opens

dorsally (Fig. 254). In others this aperture
is considerably reduced and opens more
anteriorly (Figs 258, 259). The former condi-
tion is termed Type 2A genitalia; the latter is
Type 2B. In Type 2A genitalia parameres,
volsellae and ventral setae may or may not
be present; in Type 2B genitalia these
structures never occur. In many instances
taxa with 2A genitalia have only one pair of
unarmed, small vestigial lobes apically
(Fig. 228) and it often is difficult to de-
termine if parameres or volsellae are repre-
sented; the decision often is based on
position (medial and adjacent to the midline,
or lateral). These vestigial structures are not
easily observed with the light microscope.

Distinguishing between Type 1 and the
most generalized state of Type 2 genitalia
such as occurs in the Paracentrobiini and
Tumidiclava (e.g. Figs 224-230) may cause
confusion. In both cases aedeagal apodemes
occur. However, unlike the condition in
Type 1, in Type 2 taxa the genital capsule is
indistinguishable from the aedeagus apical-
ly. Also, as indicated, in Type 2 groups the
position of the apodemes in large series of
specimens is never seen to vary relative to
the capsule, suggesting that they do not
move independently. The genitalia of Soi-
kiella may represent an intermediate state
between types 1 and 2. In this genus, the
aedeagus can be distinguished from the
capsule apically (Figs 212, 214), yet SEM
examination shows what appear to be poorly
sclerotized lateral connections (Fig. 213).

CLASSIFICATION

Several tribal classifications of the
Trichogrammatidae have been proposed
in the past century (e.g. Girault 1912, 1918;
Kryger 1918, Blood 1923, Viggiani 1971,
Yousuf and Shafee 1988). All are phenetic
and utilize few characters. Because a phy-
logenetic study of family relationships
based on molecular evidence currently is
underway (Owen et al., in prep.) this topic
is not covered in detail here. The arrange-
ment of genera below represents a modifi-
cation of Viggiani's 1971 classification, the

Journal of Hymenoptera Research

most widely followed currently. It is based aedeagal apodemes. These genera are

primarily on male genitalia. treated here as Chaetostrichini.

Viggiani divided the family into two Generic membership of the Paracentro-

subfamilies and four tribes, Trichogramma- biini (Type 2a genitalia) and Ohgositim

tinae (Trichogrammatini, Paracentrobiini) (Type 2b genitalia), with one exception

and Oligositinae (Oligositini, Chaetostri- (Ufensia), retains the criteria erf Viggiani,

chini). With exceptions, genitalic structure whereas membership of the Trichogram-

is the most generalized in the Trichogram- matini (Type 1 genitalia) and Chaetostri-

matini and the most reduced in the Oligo- chini (Type 2a genitalia) changes consider-

sitini. The other two tribes represent in- ably. The former two tribes appear to be

termediate conditions. One feature defining monophyletic, whereas the latter two are

subfamily is the presence (Trichogrammati- paraphyletic at best. The compelling reason

nae) or absence (Oligositinae) of aedeagal for deviating from Viggiani's classification

apodemes. Within the Trichogrammatinae is that the presence of aedeagal apodemes

Viggiani separated the nominate tribe from nGw is known to vary within genera (e.g. in

the Paracentrobiini primarily by genitalic Aphelinoidea, Chaetogramma, Burksiella) thus

shape and dimensions of the anterodorsal negating their value for defining tribes and

aperture. In the Trichogrammatini the gen- subfamilies. Removal of the Paracentrobiini

ital capsule is widest near the middle with from Trichogrammatinae allows recogni-

an aperture of varying shape, whereas in the non or a monophyletic Oligositinae, defined

Paracentrobiini the capsule is subconical, by the fusi0n of the aedeagus and genital

widest at its base and the aperture is capsule apically and often basally as well,

relatively short and transverse. Aperture Monophyly also is supported by the molec-

size also was used to distinguish the mar evidence (Owen et al., in prep.).

Chaetostrichini from Oligositini within the Genera are presented in the text in

Oligositinae. In the former it is relatively alphabetical order within tribe /subtribe. A

large and well defined; in the latter it is list of all trichogrammatid genera (except

considerably reduced. The genera originally fossil groups) and the subfamily and tribal

assigned to these tribes are given by assignments recognized in this work follow.

Viggiani (1971, 1984). Only one tribe, the Oligositini, is further

The arrangement followed here does not divided into subtribes. Numbers associated

purport to be strictly monophyletic. Some with genera refer to order of presentation in

arrangement of genera is necessary and the the text Genera m brackets are not known

one used hopefully is more heuristic than frQm the New WorM and are not formally

simple alphabetical presentation. It devi- treated further An asterisk (*) following

ates from Viggiani's classification in the & g name mdicates that the male is

following respects: The Trichogrammati- unknown> These genera are tentatively in-

nae consists of a single tribe and includes orated into the classification based on

all genera with Type 1 genitalia. Genera characters correlated with type of male

with Type 2 genitalia are treated as nitalia. Family-group characters and a dis-

Oligositinae. Soikiclla, with annectant gen- ° , , , , . .,,. . .u„i

,& ' _ y. cussion of generic relationships within tribal

italic structure, is placed in the Incho- .u . , . A- . . ,

. . T ' ,r. „,. ... groups are given in the text immediately

grammatmi. In addition to the Oligositini P , ,, c ,XT ,Ar ,, . , . •

6 , _ ..... „, . . . . before the first New World genus treated in

and Chaetostrichini, the Oligositinae in- . „ , ° c

, , . _, . .. . , ° , . ,, that category, ror example, the taxonomy ot

eludes the Paracentrobiini formerly in the . _ . ° J r „ . .

„ . . ,. „ , the Trichogrammatinae: lnchogrammatini

Trichogrammatinae, as well as several . ,° . _ , ° r

.Ll rr n , ,. ,, precedes the treatment or Bracnyufens.

genera with Type 2 genitalia originally r J J

treated by Viggiani as Trichogrammatini Trichogrammatinae. Trichogrammatini. [Ap-

presumably because of the presence of seudogratmna Girault]*, [Asynacta Forster]*, [Aus-

Volume 15, Number 1, 2006

tralufens Girault], [Brachyia Strand], Brachyufens
Viggiani (1), Ceratogramma De Santis (2), [Eu-
trichogramma K. Lin]*, Haeckeliania Girault (3),
[Hispidophila Viggiani], Hydrophylita Ghesquiere
(4), [Japania Girault], Mirufens Girault (5),
[Neobrachista Girault], [Neobrachistella Girault],
[Neocentrobia Girault]*, [Neocentrobiella Girault]*,
[Oligositoides Doutt], [Ophioneuris Ratzeburg],
Pachamama Owen and Pinto (6), Paratricho-
gramma Girault (7), Poropoea Forster (8), [Pro-
chaetostricha K. Lin], [Pseudogrammina Ghes-
quiere], Pterandropln/snlis Nowicky (9), [Pseudo-
mirufens Lou], Soikiclla Nowicki (10), Thanato-
gramma n. gen. (11), Trichogramma Westwood
(12), Trichogrammatella Girault (13), Trichogram-
matoidea Girault (14), Trichogrammatomyia Gir-
ault (15), [Urogramma Girault], Viggianiella n.
gen.* (16), Xenufens Girault (17), [Xenufensia
Girault], [Zelogramma Noyes and Valentine].

Oligositinae. Paracentrobiini: Ittys Girault
(18), Ittysella Pinto and Viggiani (19), Paracen-
trobia Howard (20), [Paraittys Viggiani]. Chae-
tostrichini. Adelogramma n. gen. (21), Adryas
Pinto and Owen (22), Aphelinoidea Girault (23),
Bloodiella Nowicky (24), Brachista Walker (25),
Brachygrammatella Girault (26), Burksiella De

Santis (27), Centwbiopsis Girault (28), Chacto-
gramma Doutt (29), Chaetostricha Walker (30),
[Densufens N. Lin], Lathromeris Forster (31),
[Kyuwia Pinto and George], Lathromeroidea Gir-
ault (32), [Lathromeromyia Girault], Monortho-
chaeta Blood (33), Nicolavespa Pinto (34), [Parus-
canoidea Girault], Pintoa Viggiani (35), [Pseudo-
brachysticha Girault]*, Psenduscana n. gen. (36),
Pteranomalogranima Viggiani and Velasquez*
(37), Pterygogramma Perkins (38), [Thoreauia
Girault]*, Tumidiclava Girault (39), Tumidifemur
Girault (40), Ufens Girault (41), [Ufensia Girault],
Uscana Girault (42), Uscanella Girault* (43),
Uscanoidea Girault (44), Uscanopsis Girault (45),
Xiphogramma Nowicki (46), Zaga Girault (47),
Zagella Girault (48). Oligositini. Oligositina:
Epoligosita Girault (49), Megaphragma Timberlake
(50), Oligosita Walker (51), Prestwichia Lubbock
(52), [Prosoligosita Hayat and Hussein], Sinepalpi-
gramma Viggiani and Pinto* (53). Eteroligositina:
[Chaetostrichella Girault], Doirania Waterson (54),
[Eteroligosita Viggiani], [Hayatia Viggiani], [Pro-
brachista Viggiani]*, Pseudoligosita Girault (55).

Genus unplaced to subfamily. [Neola-
thromera Ishii]*

KEY TO THE NEW WORLD GENERA OF TRICHOGRAMMATIDAE
(Numbers preceeding genera in key refer to order of presentation in text)

1. Scutellum with at least 2 pair of setae (anterior pair may be very short), midlobe of
mesoscutum usually with 2 or more pair of setae (Figs 95, 179, 180, 184, 185). Eyes
light to dark red in color 2

1'. Scutellum and midlobe of mesoscutum each with 1 pair of setae (Figs 186, 187). Eyes

black. (Oligositini) 60

2(1 ). Fore wing very narrow, 5-10X as long as wide (Figs 114a, b). Antenna with 2 funicular
segments, both longer than wide; female club with two large spatulate sensilla
(Fig. 15) 4. Hydrophylita

2'. Fore wing broader, considerably less than 5X as long as wide. Antenna with funicle

variable in structure or entirely absent; female club without large spatulate sensilla. 3

3(2'). Antenna with 1 funicular segment (segment may be partially divided near middle as

in Fig. 49) 4

3'. Antenna either with funicle absent or consisting of 2 segments (Fl often small and

closely appressed to base of F2 as in Fig. 5) 8

4(3). Antennal club 1 segmented

4'. Antennal club 2-3 segmented 6

5(4). Fore wing densely setose (Fig.126); marginal vein (MV) contacting anterior margin of
wing. Funicular segment partially divided (Figs 48, 49). Mesophragma notched
apically (Fig. 193). Petiolar segment (immediately behind propodeum) without
a transverse row of denticles 29. Chaetogramma (in part)

s2 Journal of Hymenoptera Research

5' Fore wing sparsely setose (Fig. 110); MV not attaining anterior margin of wing.
Funicular segment strongly constricted apically (Fig. 28) but not partially divided
into 2 segments. Mesophragma entire apically (Fig. 194). Petiolar segment with
a transverse row of denticles (Figs 188, 189) 7. Paratrichogramma 9

6(4')- Fore wing narrow, 2.5-3.5 X as long as wide with elongate fringe setae at least half fore
wing width (FWW) (Fig. 129). Funicle with a strongly curved J-shaped placoid
sensillum (PLS) (Fig. 52B). Midlobe of mesoscutum with 1 or 2 pair of setae;
scutellum with anterior pair of setae much shorter than posterior pair (Figs 184-
•■nri 35. Pintoa

6'. Fore wing wider, c. 1.5X as long as wide, with fringe setae short, much less than half

FWW (Figs 111, 148). Funicle with PLS not strongly curved (Fig. 30). Both midlobe
of mesoscutum and scutellum each with two pair of elongate setae (Fig. 180). ... 7

7(6'). Fore wing without an RSI setal track (Fig. 111). Petiolar segment with a transverse row

of denticles (Figs 188, 189) 10- Soikiella 9

7'. Fore wing with a well developed RSI setal track (Fig. 148). Petiolar segment without

a transverse row of denticles 24. Bloodiella

8(3'). Hind tibial spur extremely long and truncate apically, c. 0.9 x the length of entire hind

tarsus 45- Uscanopsis

8'. Hind tibial spur considerably shorter, pointed apically, at most slightly longer than
first tarsomere

9(8'). Antenna with only 1 elongate postanellar segment (Fig. 35)

12. Trichogramma (Trichogmmma) 6*

9'. Antenna with more than 1 postanellar segment 10

10(9'). Fore wing venation strongly sinuate with greatest curvature basally, between
premarginal and marginal veins; posterior margin of wing with a distinct
preretinacular lobe (Fig. 103). Male scape enormously inflated, its width greater
than the length of club (Fig. 19); female scape unmodified (Fig. 18). . . 6. Pachamama

1 0'. Fore wing venation not as above, either not sinuate at all, or if so, then apical curvature
similar to or greater than basal curvature (Figs 104, 113); posterior margin of wing
without a preretinacular lobe. Male scape at most slightly inflated, its width never
approaching length of club H

11(10'). Antenna with a 2-segmented funicle and a 1-segmented club 12

11'. Antenna with funicle present or not but club at least 2 segmented 19

12(11). Fore wing with abbreviated venation, stigmal vein (SV) reduced to a small appendix at
apex of marginal vein (MV) (Fig. 110). Antenna with second funicular segment (F2)
noticeably narrowed at apex, bottle shaped (Fig. 29) 7. Paratrichogramma <$

12'. Fore wing venation not so abbreviated, SV variable but clearly diverging from MV,
never reduced to a small appendix. Antenna with F2 not narrowed at apex, not
bottle shaped 13

13(12'). Fore wing (Fig. 116) with stigmal vein (SV) more than half the length of marginal vein
(MV), positioned parallel to anterior wing margin, reaching 0.8 the wing length
(FWL); fore wing disk virtually glabrous. Hind wing disk also glabrous, with

anterior fringe very long, almost as long as posterior fringe (Fig. 98)

16. Viggianiella 9 (J1 unknown)

13'. Fore wing with SV variable in length but distinctly divergent from anterior wing
margin, not reaching beyond 0.6 FWL; fore wing disk noticeably setose. Hind wing
disk with one or more setal tracks, its anterior fringe considerably shorter than
posterior fringe 14

Volume 15, Number 1, 2006 53

14(13'). Fore wing venation sinuate, with marginal vein (MV) gradually curving away from
anterior wing margin onto stigmal vein (SV) (Figs 113, 114). Mesophragma entire
apically (Fig. 194) 15

14'. Fore wing venation not sinuate; MV ending abruptly distally, not gradually curving

onto SV (Fig. 115). Mesophragma notched apically (Fig. 193) 16

15(14). Fore wing with RSI track present behind stigmal vein (SV) (Figs 113, 158); premarginal
vein (PM) with 2 setae. Hind wing usually with at least 2 setal tracks (a complete
middle track and a complete or partial posterior track) 12. Trickogramma 9

15'. Fore wing without an RSI track behind SV (Fig. 114); PM with 1 seta. Hind wing with

only a middle setal track which is not complete to wing apex

14. Trichogrammatoidea Q

16(14'). Fore wing (Fig. 115) fringe setae elongate, their maximum length at least half wing
width (FWW); venation evenly sclerotized; fore wing disk only moderately densely
setose, setae arranged in linear tracks which continue to apex of wing. Antenna 1
club unevenly tapering to apex (Fig. 37) 15. Trichogrammatomyia o (in part)

16'. Fore wing (Figs 125, 126) fringe setae short, their maximum length no more than 0.2 X
FWW; venation unevenly sclerotized, base of marginal vein and apex of
premarginal vein much lighter in coloration; fore wing disk densely setose, setae
scattered, linear setal tracks poorly indicated especially in apical fourth of wing.
Antennal club evenly tapering to apex (Fig. 48) 17

17(16'). Ovipositor elongate, extending entire length of metasoma and well beyond its apex
(Fig. 195). Funicle with 2 completely divided segments (Fig. 62). Male without
aedeagal apodemes (AAP) (Fig. 239) 46. Xiphogramma

17'. Ovipositor shorter, not extending entire length of metasoma, nor appreciably beyond
its apex. Funicle either distinctly 2 segmented or only partially divided. Male with
AAP (Fig. 229) 18

18(17'). Fore wing with marginal vein (MV) densely setose (Fig. 125); funicle distinctly two

segmented 26. Brachygrammatella (in part)

18'. Fore wing with MV not densely setose (Fig. 126); funicle only partially divided

(Fig. 49) 29. Chaetogramma (in part)

19(11'). Maxillary palp 2 segmented (Fig. 173) 20

19'. Maxillary palp 1 segmented (Fig. 172) 27

20(19). Antenna with a distinct 2-segmented funicle 21

20'. Antenna without a funicle 25

21(20). Fore wing with marginal vein (MV) extremely short and broad, subquadrate, much

shorter and wider than premarginal vein (PM) (Fig. 108). ... 1. Brachyufens (in part)

21'. Fore wing with MV not subquadrate, distinctly longer than wide, its length relative to

that of PM variable, but never much wider 22

22(21'). Pedicel of antenna with transverse, crenulate ridges (Fig. 16); male club with a small
fourth segment (Fig. 17). Fore tibia with several thorn-like spines on dorsal surface
(Fig. 178) 5. Mirufens

17! . Pedicel of antenna relatively smooth, without transverse, crenulate ridges; male club
usually 3, less commonly 4 segmented. Fore tibia lacking thorn-like spines on
dorsal surface (Fig. 175) 23

54 Journal of Hymenoptera Research

23(22'). Fore wing venation distinctly sinuate; stigmal vein (SV) perpendicular to marginal

vein (MV) (Fig. 104) 8. Poropoea

23'. Fore wing venation not sinuate; SV slanted diagonally away from MV 24

24(23'). Antenna with 2 anelli; first funicular segment (Fl) strongly transverse (Figs 22, 23);
male antennal club 3 segmented. Male fore wing venation enormously inflated,
occupying almost entire basal width of wing (Fig. 106). Male genitalia without an
intervolsellar process (IVP) (Fig. 207) 9. Pterandrophysalis

24'. Antenna with 3 anelli (Fig. 1); Fl usually longer than wide or subquadrate, rarely
somewhat transverse; male antennal club 3 or 4 segmented. Male fore wing
venation not inflated (Figs 99, 100). Male genitalia with an elongate IVP (Figs. 201,
202) 2. Ceratogramma

25(20'). Fore wing venation sinuate, its greatest curvature apically where marginal vein (MV)
gradually curves away from anterior wing margin onto an elongate, slender
stigmal vein (SV) (Fig. 107). Petiolar segment without a transverse row of
denticles 13. Trichogrammatella

25'. Fore wing venation not sinuate; MV ending abruptly distally, not gradually curving
onto SV (Figs 108, 109). Petiolar segment with a transverse row of denticles
(Figs 188, 189) 26

26(25'). Midlobe of mesoscutum with 3 pair of elongate setae (Fig. 179). Fore wing with
marginal vein (MV) short, subquadrate, distinctly shorter and c. 3X wider than
premarginal vein (PM) (Fig. 108) 1. Brachyufens (in part)

26'. Midlobe of mesoscutum with 2 pair of elongate setae (Fig. 180). Fore wing with MV
not subquadrate, its width considerably less than 3X that of PM (Fig. 109). . . .
3. Haeckeliania

27(19'). Antenna with funicle present 28

27'. Antenna with funicle absent 44

28(27). Fore wing venation sinuate, greatest curvature distally where marginal vein (MV)
gradually curves away from anterior wing margin onto stigmal vein (SV) (Figs 113,
114, 117) 29

28'. Fore wing venation not sinuate, MV ending abruptly distally, not gradually curving

onto SV (as in Figs 128, 135, 149) 31

29(28). Antennal club distinctly 3 segmented; funicular segments subquadrate or only slightly
transverse (Fig. 36). Fore wing with marginal vein (MV) attaining anterior margin
of wing 30

29'. Antennal club with only 2 complete segments, a third (if present) incompletely
separated from second; funicular segments strongly transverse (Figs 40-42). Fore
wing with MV placed slightly behind anterior margin of wing 17. Xemifens

30(29). Fore wing with RSI present (Fig. 113); premarginal vein (PM) with 2 setae. Male

genitalia with a dorsal lamina (DLA) (Figs. 198, 218)

12. Trichogramma (Vanlisus) $

30'. Fore wing without an RSI (Fig. 114); MV with only 1 seta. Male genitalia without

a DLA (Fig. 220) 14. Trichogrammatoidea 6*

31(28'). Antenna with the first funicular segment (Fl) distinctly shorter and narrower than the

second (F2) and closely appressed to F2 (Figs 51, 71, 84, 267, 268) 32

31'. Antenna with Fl as wide as F2 and usually subequal to or longer than F2 (Figs 31-33,
43-45, 59, 60), if shorter (Fig. 38) then dorsum of petiolar segment with a transverse
row of denticles (Figs 188, 189) 36

Volume 15, Number 1, 2006 55

32(31). Female antennal club with a terminal process (Fig. 51). Male genitalia with apodemes
(AAP) of aedeagus and volsellae (VS) present (Figs 232, 233). North and Central
America 34. Nicolavespa

32'. Female antennal club without a terminal process. Male genitalia with AAP and VS

usually absent, if present then occurring in South America 33

33(32'). Fore wing unique, almond shaped, apical margin pointed; disk without setae behind
venation; RSI absent; fringe setae elongate, their length at least half the fore wing
width (FWW) 37. Pteranomalogramma (5 unknown)

33'. Fore wing not almond shaped, apical margin broadly rounded; disk with setation
behind venation; RSI present; fringe setae shorter, their length never approaching
half FWW 341

34(33'). Fore wing venation (Fig. 150) without a constriction between stigma and marginal vein
(MV); stigma not extending apicad to apex of MV (a line drawn through stigmal
vein to wing margin describes a right angle with MV); RSI usually of 5 or fewer
setae, arranged in a straight line, directed toward posterior margin of wing,
subperpendicular to Cu tracks. Male genitalia with ventral setae (Fig. 255). . . .
48. Zagella

34'. Fore wing venation (Figs 142, 143) with a constriction between stigma and MV; stigma
extending apicad to apex of MV (a line drawn through stigma to wing margin
describes an oblique angle with MV); RSI longer, consisting of more than 5 setae,
distinctly curved or arranged in a straight line but directed toward base of wing
and usually converging with Cu tracks. Male genitalia almost always without
ventral setae (Figs 243, 245) 35

35(34'). Antenna with funicle narrower than club (Fig. 72A). Fore wing with RSI relatively
straight (Fig. 143). Foretibia with a prominent thorn-like spine at middle of dorsal
surface (Fig. 176). Ovipositor usually extending beyond apex of metasoma. Male
genitalia narrow and elongate, almost always as long or longer than hind tibia
(Figs 244, 245). North America 30. Chaetoshicha

35'. Antenna with funicle slightly wider than club (Fig. 69). Fore wing with RSI broadly
curved (Fig. 142). Fore tibia without a thorn-like spine at middle of dorsal surface.
Ovipositor shorter, not extending beyond apex of metasoma. Male genitalia short,
broader (Figs 242, 243), considerably shorter than hind tibia. Broadly distribut-
ed 27. Burksiella

36(31'). Fore wing with long fringe setae, their maximum length at least half width of wing
(FWW) (Fig. 115). Antenna with funicular segments distinct, Fl shorter or equal in
length to F2 (Figs 37, 38) 15. Trichogrammatomyia o (in part), ;

36'. Fore wing usually with fringe setae distinctly shorter than half FWW but if longer then

funicle with F2 noticeably shorter than Fl 37

37(36'). Antenna with both funicular segments, and first 2 club segments (CI, C2) each with
several thick, spiniform placoid sensilla (PLS) attached to antennal surface only at
base (Fig. 31) 10. Soikiella ;

37'. Antenna without thick spiniform PLS; funicle either without PLS entirely (Figs 32, 33,

43-45) or with all but apex of PLS attached to surface (Figs 59, 60) 38

38(37'). Antenna with funicular segments each with at least 1 placoid sensillum (PLS); funicle

broad, subequal in width to base of club 39

'An apparent new genus, very recently collected from several localities in central Chile, keys to this point (RSI absent,
however) but fits neither part of couplet 34. Only females are known. Its antenna is typical or certain members ot the
Chaetostricha group such as Cbaetostricha and Zagella (2-segmented funicle, Fl very short and appressed to F2, club 3
segmented) but the fore wing is unique. The apparent absence of a stigmal vein and the lack ot the RSI track separate it from
other genera with similar antennal structure. Actually a stigmal vein is present in this taxon but it lacks all pigmentation.

i-. Journal of Hymenoptera Research

38'. Antenna with funicular segments lacking PLS; funicle distinctly narrower than base of

club (Figs 32, 33, 43-45) 41

39(38). Fore wing (Fig. 135) moderately densely setose with distinct linear setal tracks
throughout, including a well defined RSI. Male antennal club 4 segmented
(Fig. 60) 4L Ufms

39'. Fore wing densely setose, without well defined linear setal tracks, RSI absent. Male

antennal club with fewer than 4 segments 40

40(39'). Fore wing with marginal vein (MV) densely setose, with more than 10 setae dorsally
(Fig. 125); stigmal vein (SV) indistinct, short, without a constriction between stigma
and MV 26- Bmchygrammatella (in part)

40'. Fore wing with MV normally setose, with fewer than 10 setae dorsally; SV distinct,

with a constriction between stigma and MV 33. Monorthochaeta

41(38'). Petiolar segment with a transverse row of denticles dorsally (Figs 188, 189). Fore wing
(Fig. 112) without an RSI; fringe setae short, c. 0.1 wing width (FWW). Female
antennal club (Fig. 32) 2 segmented, apical segment (C2) with a patch of recurved
setae (RS) ventroapically; male antennal club 3 segmented. Male genitalia with
aedeagus (ADG) distinct from genital capsule (GC) and much longer than capsule
(Figs 215-217) 11- Thanatogramma

41'. Petiolar segment without a transverse row of denticles dorsally. Fore wing with or
without an RSI; fringe setae variable but length greater than 0.1 FWW. Antennal
segment number similar in both sexes. Female club without a patch of RS
ventroapically on apical segment. Male genitalia with ADG incorporated into GC
(Figs 224-226) 42

42(41'). Antennal club 2 or incompletely 3 segmented; only 1 anellar segment present (Fig. 45).

Fore wing with RSI absent (Fig. 120) 19. Utysella

42'. Antennal club distinctly 3 segmented; 2 anelli present (Figs 43, 44). Fore wing with RSI

present or absent (Figs 118, 119) 43

43(42'). Funicular segments relatively elongate (Fig. 44), the first (Fl) longer than wide, second
(F2) about as long as wide. Fore wing with RSI present (Fig. 119). Male with
sternum VIII modified into two broad plate-like appendages (Fig. 225). ... 18. Htys

43'. Funicular segments shorter (Fig. 43), Fl about as wide as long, F2 transverse, distinctly

broader than long. Fore wing usually without a distinct RSI (Fig. 118). Male

sternum VIII modified into two narrow, strap-like appendages (Fig. 224)

20. Paracentrobia

44(27'). Head with a unique deep and relatively large pit immediately in front of anterior
ocellus (Figs 165-167). Metasomal tergum II (first visible) with an oval, clear bulla
anterolaterally (Fig. 196) 32. Lathromeroidea (in part)

44'. Head without a deep pit in front of anterior ocellus. Metasomal tergum II (first visible)

lacking anterolateral bullae 45

45(44'). Fore wing with stigmal vein (SV) short, sessile, not distinct from marginal vein (MV);

disk densely setose, RSI not indicated. Figs 121-123, 159 46

45'. Fore wing with a relatively elongate SV which is distinct from MV; disk not densely

setose, RSI usually present. As in Figs 140, 144, 149 47

46(45). Female antennal club with a distinct terminal process at apex (Fig. 56). Male
metasomal tergum VII (and often VI as well) with a pair of lateral suboval or
reniform pustules (as in Fig. 264). Fore wing with setal tracks R and RS2 distinct and
well separated (Fig. 132). Hind wing with only 2 complete setal tracks, the third
(posterior) track absent at least in basal half 39. Tumidiclava

Volume 15, Number 1, 2006 57

46'. Female antennal club without a terminal process at apex (Fig. 46). Male metasomal
terga without pustules. Fore wing disk more uniformly and densely setose, without
distinct R and RS2 setal tracks (Figs 121-123). Hind wing with at least 3 complete
setal tracks 23. Aphelinoidea

47(45'). Antennal club with first segment (CI) very short, transverse, usually closely appressed
to base of second (C2), without placoid sensilla (PLS), its width not approaching
maximum club width (Figs 6, 65, 66, 73, 74, 76-83) 48

47'. Antennal club with CI longer, variable in shape but its width subequal to maximum

club width, often bearing PLS (Figs 58, 61) 54

48(47). Fore wing of female (Fig. 140) with premarginal (PM) and marginal veins (MV)
confluent; PM subtriangular, widest basally, distinctly wider than MV and with
a distinct gap between subcostal (SC) and PM veins. Fore wing of male similar or,
in some species, with PM and base of MV inflated and subequal in width
(Fig. 141) 22. Adryas

48'. Fore wing not as above. PM not subtriangular in shape, narrower than or subequal in
width to MV; PM and MV confluent or not. Fore wing of male never with inflated
venation 49

49(48'). Fore wing with RSI absent; disk sparsely setose overall, subglabrous between r-m and

Cu2; fringe setae elongate, length c. half width of wing (FWW)

43. Uscanella Q ( ^unknown)

49'. Fore wing with RSI present; disk moderately densely setose throughout; fringe setae

varying in length but usually considerably less than half FWW 50

50(49'). Fore wing venation (Figs 149, 162) without a constriction between stigma and
marginal vein (MV); stigma not extending apicad to apex of MV (a line drawn
through stigmal vein to wing margin describes a right angle with MV). Foretibia
usually with a prominent thorn-like spine at middle of dorsal surface
(Fig. 177) 47. Zaga

50'. Fore wing venation (Figs 143, 144, 147, 161) with a constriction between stigma and
MV; stigma extending apicad of apex of MV (a line drawn through stigmal vein to
wing margin describes an oblique angle with MV) (rarely without a well defined
stigmal vein). Fore tibia variable, rarely spinose but if so without a more prominent
spine at middle of dorsal surface (Figs 175, 178) 51

51(50'). Female antennal club obsolescently segmented (Figs 63, 64, 66), only 3 complete
segments present: a minute CI, a short transverse C2, and an elongate and only
partially divided C3/4/5; with at least 1 placoid sensillum (PLS) spiniform (free of
surface entire length) and with PLS on C2 extending beyond surface for at least half
its length. Male antenna unique (Fig. 65), club distinctly 5 segmented, C3, C4 and
C5 each with 1 or more long spiniform PLS which exceed the length of the club
itself 21. Adelogramma

51'. Male and female antennal club both 5 segmented (Figs. 73-78); PLS not spiniform, not

extending far beyond antennal surface 52

52(51'). Ovipositor elongate, extending entire length of metasoma with c. V* of its length
extending beyond metasomal apex. Male genitalia elongate, narrow, longer than
hind tibia (HTL), with a longitudinal furrow ventrally (Fig. 251). . . 28. Centrobiopsis

52'. Ovipositor shorter, not extending appreciably beyond apex of metasoma. Male
genitalia considerably shorter with length less than HTL and without a longitudinal
furrow ventrally (Figs 249, 250) 53

53(52'). Propodeal disk and metanotum usually subequal in length (Fig. 182). Fore wing
relatively broad (usually < 2X as long as wide) with short fringe setae (usually
< 0.2 wing width) and a broadly curved RSI (Fig. 147). Male genitalia usually
without ventral setae (Fig. 250) 44. Uscanoidea

j-e Journal of Hymenoptera Research

53'. Propodeal disk longer than metanotum (Fig. 181). Fore wing narrower (> 2X as long
as wide) with longer fringe setae (> 0.2 wing width); RSI, if obvious, usually
relatively straight, not broadly curved (Figs 144-146). Male genitalia usually with
ventral setae (Fig. 249) 32. Lathromeroidea (in major part)

54(47'). Antennal club 2-4 segmented 55

54'. Antennal club 5 segmented

55(54). Fore wing disk moderately to densely setose, without distinct setal lines at least in

posterior two-thirds; RSI usually absent (Figs 124, 133, 134) 56

55'. Fore wing disk moderately densely setose, with distinct setal lines including RSI

(Figs 130, 136) 57

56(55). Fore wing with several short stiff setae in costal cell and on marginal vein (MV);
stigmal vein (SV) diverging only slightly from MV (Figs 133, 134, 160). Female
without strut-like apodemes extending anteriorly from near base of oviposi-
ts 40. Tumidifemur

56'. Fore wing without numerous setae in costal cell and on MV; SV strongly divergent
from MV (Fig. 124). Female with a pair of strut-like apodemes extending anteriorly
from near base of ovipositor (Fig. 265) 25. Brachista

57(55'). Antennal club with first segment (CI) as long or longer than second (C2); female with
1 or more placoid sensilla (PLS) on CI (Fig. 61). Uniformly brown in color. Male
genitalia simple, without volsellae (VS) or parameres (PAR) (as in Fig. 250). . .

42. Uscana

57'. Antennal club with CI distinctly shorter than C2, CI without placoid sensilla in both
sexes (Figs 53, 54). Distinctly bicolored, brown with white coloration on much of
mesosoma and legs. Male genitalia variable, most species with distinct VS and PAR
(as in Fig. 235) 36. Pseuduscana

58(54'). Female antennal club with a terminal process at apex (Fig. 50). Fore wing with RSI
absent, discal setation moderately dense at most (Fig. 127). Male metasomal tergum

VI (and often VII) with modified areas as indicated in Figs 262-264

31. Lathromeris

58'. Female antennal club without a terminal process at apex. Fore wing with an RSI
(Figs 131, 144, 145), if RSI not obvious then disc densely setose, obscuring most
linear setal tracks (Fig. 146). Male metasomal terga without modified areas 59

59 (58'). Antennal club with first segment (CI) longer than second (C2); C2 transverse, shortest
segment of club; CI with a placoid sensillum (PLS) (Figs 55, 94). Hind wing disk
usually with only 1 setal track. Male genitalia with aedeagal apodemes (AAP)
(Fig. 96) 38. Pterygogramma

59'. Antennal club with CI distinctly shorter than C2; CI shortest segment of club and
without a PLS (Figs 73-77). Hind wing disk with at least 2 linear setal tracks. Male
genitalia without AAP (Figs 246, 248) 32. Lathromeroidea (in part)

60(1'). Maxillary palp entirely absent (Fig. 171). Antenna without placoid sensilla (PLS) on

surface. Mesosomal dorsum exceedingly smooth, without reticulae

53. Sinepalpigramma

60'. Maxillary palp present (Fig. 172). Antenna with PLS on surface of club. Mesosomal

dorsum sculptured, at least with obsolescent reticulae 61

61(60'). Fore wings very narrow, c. 7x as long as wide (Figs 152, 154) 62

61'. Fore wings broader, no more than 4X as long as wide (Figs 151, 153, 156, 157). ... 63

62(61). Antenna with 3 postanellar segments (Fig. 87). Fore wing sparsely setose with only 1

or 2 rows of setae on disk at most (Fig. 152) 50. Megaphragma

Volume 15, Number 1, 2006

62'. Antenna with 4 postanellar segments (Fig. 90). Fore wing densely setose

(Fig. 154) 52. Prestwkhia

63(61'). Metasomal terga uniformly sclerotized their entire length. Propodeal disk usually
distinctly longer than metanotum at midline (Fig. 186). Mesopleuron with pleural
suture (Fig. 192) 64

63'. Metasoma with at least the 3 anterior terga longitudinally striate posteriorly,
uniformly sclerotized anteriorly (Figs 187, 197). Propodeal disk only slightly longer
than metanotum (Fig. 187). Mesopleuron without a pleural suture (as in Figs 191,
197) 65

64(63). Fore wing disk entirely glabrous (with at most 1 or 2 setae) (Fig. 151). Antennal club 1
or incompletely 2 segmented. Mesosoma with a small subtriangular lobe arising
beneath propodeal disk. Mesotarsus elongate, distinctly longer than mesoti-
bia 49. Epoligosita

64'. Fore wing disk sparsely to moderately densely setose (Fig. 153). Antennal club 3
segmented. Mesosoma without a subtriangular lobe beneath propodeal disk.
Mesotarsus not longer than mesotibia 51. Oligosita

65(63'). Antennal club 1 segmented (Fig. 92). Male genitalia simple, only slightly curved

ventrally, without posteriorly directed apodemes at base (as in Fig. 258)

54. Doirania

65'. Antennal club 3 segmented (Fig. 93). Male genitalia strongly curved ventrally, with

posteriorly directed apodemes at base (Fig. 259) 55. Pseudoligosita

GENERIC REVIEWS ARRANGED BY
SUBFAMILY AND TRIBE

Trichogrammatinae

Diagnosis. — Maxillary palp 1 or 2 segment-
ed. Male. Type 1 genitalia. GC and ADG dis-
tinct, ADG capable of movement indepen-
dently of GC and discernible from GC its entire
length; GC with VS and PAR (apparently unit-
ed into a single lobe in Xenufens). Figs 198-223.

Discussion. — The Trichogrammatinae
consists of the nominate tribe only. Its
definition differs from that of Viggiani
(1971) by excluding genera having male
genitalia in which the aedeagus and genital
capsule are consolidated and indistin-
guishable apically but with aedeagal apo-
demes basally (Figs 224-230). Excluded
from the subfamily on this basis are the
Paracentrobiini as well as certain genera
previously placed in the Trichogrammatini
(Aphelinoidea, Brachygrammatella, Lathro-
meris, Monorthochaeta, Tumidiclava, Ufens).
All are herein assigned to the Oligositinae,
the latter six genera to the Chaetstrichini.

Trichogrammatini Westwood

Discussion. — Most genera of Trichogram-
matini can be assigned to one of two
phenetic groups. In one (Group 1) the
antenna has five postanellar segments in
both sexes, the maxillary palp usually is
two segmented, axillae and axillulae are
relatively well separated from the scutel-
lum, and the male genital capsule is open
its entire length dorsally and trough-like in
appearance. These are generalized fea-
tures. The male genitalia retain several of
the basic features characterizing Hyme-
noptera (Snodgrass 1957), the antennae are
minimally dimorphic sexually with seven
to nine flagellomeres (the maximum num-
ber for the family), consolidation of the
mesosomal tergites is minimal, and the
maxillary palp remain unreduced (nomi-
nate subgenus of Hydrophylita an excep-
tion). New World genera with these char-
acteristics include Brachyufens, Cerato-
gramma, Haeckeliania, Hydrophylita, Miru-
fens, Poropoca, Pterandrophysalis, and

60 Journal of Hymenoptera Research

Trichogrammatella. Several extralimital gen- oval club of one or two closely appressed
era such as Ophioneurus and Oligositoides segments with short funicular segments
also belong here. I am aware of no derived lacking placoid sensilla. Four of these
feature shared by these taxa and this basal genera (Thanatogramma, Trichogramma, Tri-
assemblage is apparently paraphyletic. The chogrammatoidea, and Xenufens) have mand-
majority of genera in this group with known ibles with a large socketed apical tooth, as
hosts are parasitoids of Curculionoidea. far as I know the only occurrence of this
Included are Brachyufens, Ceratogramma, structure in the family. Additionally, all
Haeckeliania, Ophioneurus, Poropoea, and taxa in this assemblage with known hosts
Pterandrophysalis. Coleoptera parasitization (Australufens, Paratrichogramma, Tricho-
is not common in other segments of the gramma, Trichogrammatoidea, Trichogramma-
family, suggesting that this association is tomyia, Xenufens) parasitize eggs of Lepi-
primitive for the Trichogrammatidae. Mir- doptera, an uncommon host group in other
ufens, Trichogrammatella and the African segments of the family,
genus Oligositoides parasitize eggs of auche- The row of denticles on metasomal
norrhynchous Hemiptera, hosts utilized by tergum I is a unique feature occurring only
perhaps the majority of Oligositinae. in the Trichogrammatini. There is not yet
Genera in the second group of Tricho- convincing evidence that this trait delimits
grammatini (Group 2) have more derived a monophyletic group. It occurs in all
features. The antenna has fewer than five Group 2 genera except Viggianiella and
postanellar segments at least in the female, Pachamama. It also is present in Group 1
the maxillary palp usually is one segment- genera Haeckeliania and Brachyufens.
ed, axillae and axillulae are poorly de-
lineated from the scutellum, and the first 1- Brachyufens Viggiani
metasomal (petiolar) tergum usually has (rigs 24, 25, 1U8, 179)

a transverse row of denticles. The male Bmchyufens viggiani, in Doutt and Viggiani

genital capsule is variable. It can be 1%8 Type species: ufms osborni Dozier, by

completely open dorsally and trough-like original designation,
as in Group 1 or possess a transverse

bridge dorsal to the aedeagus in the apical Diagnosis. — Female. Antenna with 2 anelli

half of the capsule. New World genera in and 5 club segments; however 3 basal-most

Group 2 include Pachamama, Paratricho- postanellar segments relatively large,

gramma, Soikiella, Thanatogramma, Tricho- loosely associated and often not easily

gramma, Trichogrammatoidea, Trichogramma- categorized as funicular or club segments;

tomyia, Viggianiella and Xenufens. The Aus- 2 terminal club segments closely appressed

tralian genera Australufens, Brachyia, Pseu- with apical-most segment subcorneal, with

dogrammina, and Urogramma also belong RS on ventral surface; postanellar segments

here. Based on male genitalia and with PLS extending considerably beyond

female antennal structure (Noyes and surface but attached for at least half their

Valentine 1989), the New Zealand genus length. Maxillary palp 2 segmented. Mid-

Zelogramma is a possible member of this lobe of mesoscutum with 3 pair of elongate

group as well. setae. Propodeum narrowing to middle,

Paratrichogramma, Thanatogramma, Tri- anterior margin broadly V-shaped. Fore

chogramma, Trichogrammatoidea, Trichogram- wing broad, oblate, less than 2X as long as

matomyia and Xenufens, as well as the four wide, fringe setae short; venation unique:-

Australian genera mentioned share several MV very short, subquadrate, distinctly

derived features. In all, the petiolar tergum shorter and c. 3x wider than PM, SV and

has a transverse row of denticles, and the radial process well developed, costal cell

female antenna is characterized by a sub- large; disk moderately densely setose with

Volume 15, Number 1, 2006

few linear setal tracks apparent, RSI
present. Metasomal tergum I (petiole) with
a transverse row of denticles.

Male. Antenna with 5 relatively large,
loosely articulated segments (5-segmented
:lub), each segment with a whorl of spini-
form PLS which are attached to surface only
at base. Genitalia similar to Haeckelimiia.

Distribution. — North America.

Diversity. — Brachyufens osborni, known
from the West Indies and southeastern
United States, is the only species described.
Originally placed in Ufeus, it was removed
and treated as the type species of Br achy u-
cens by Viggiani (in Doutt and Viggiani
1968). One or two undescribed species
3ccur in Mexico and the southwestern
United States.

Discussion. — Brachyufens is likely a de-
rived element of Haeckeliania. General body
ihape, antennal structure in both sexes, the
maxillary palpi, and male genitalia are
similar in both. Although the large quad-
rate marginal vein separates Brachyufens
from the vast majority of Haeckeliania (cf.
Figs 108, 109), there are undescribed
Australian species of the latter which
approach Brachyufens in this regard. The
number of setae on the mesoscutum also
should separate the two genera as current-
ly defined. Both genera are known to
Darasitize curculionid eggs.

Brachyufens females were characterized
i>y their two-segmented funicle and three-
segmented club in Doutt and Viggiani
^1968). Because of the loose association of
the first three postanellar segments the
antenna is difficult to characterize especial-
ly on slide-prepared material. Specimens
may appear as 2F/3C, 3F/2C or 5C depend-
ing on the preparation. The five postanellar
segments in males (previously unknown)
are all loosely associated and more clearly
:omprise a five-segmented club.

Records. — Mexico: Baja California Sur,
Veracruz. United States: California, Flor-
ida, Texas. West Indies.

Hosts. — Coleoptera: Curculionidae (Sch-
auff 1987).

2. Ceratogramma De Santis, renewed
status

(Figs 1, 3, 12-14, 99, 100, 190, 200-202)

Ceratogramma De Santis 1957. Type species:
Ceratogramma schachovskoyi De Santis, by
original designation. Pinto and Viggiani
1991 (generic review). Fursov 1995a (as
synonym of Szelenyia Nowicki).

Diagnosis. — Female. Antenna with 3 an-
elli, 2 funicular and 3 club segments;
funicular segments distinctly separate
from one another. Maxillary palp 2 seg-
mented. Midlobe of mesoscutum and
scutellum each with 2 (sometimes 3 in
one species) pair of elongate setae. Fore
wing 1. 8-2.0 X as long as wide; venation
with MV diverging from wing margin at
base, usually confluent with PM, PM
(and base of MV) strongly divergent from
wing margin, resulting in a large, often
highly setose costal cell, a short postmar-
ginal vein present or not, SV distinct,
constricted between stigma and MV,
slanted diagonally toward middle of apical
margin of wing; disk densely setose,
distinct setal tracks absent or few in
number, RSI absent.

Male. Antennal club 3-4 segmented. GC
entirely open dorsally, with an elongate
IVP which usually extends to or beyond
apex of PAR and VS.

Distribution. — Southwestern United
States, Florida (introduced), West Indies,
Central and South America in the New
World; Europe.

Diversity. — Nine species are assigned to
Ceratogramma. Eight occur in the New
World. This includes C. jeffersi, a new
species described below, the only indige-
nous Ceratogramma from the United States.
The only other species known from the
USA, Ceratogramma etiennei Delvare, was
recently introduced into Florida from the
West Indies for control of the root weevil,
Diaprepes abbreviatus (L.) (Hall et al. 2001).
The remaining six New World Cerato-
gramma include C. brasiliense Viggiani, C.
magnificum Pinto and Viggiani, C. masneri

Journal of Hymenoptera Research

Pinto and Viggiani, C. robustum Pinto and
C. schachovskoyi.

Discussion. — Ceratogramma is most simi-
lar to Pterandrophysalis and Mirufens. It is
separated from these and all other New
World genera by the three anellar seg-
ments. The postanellar antennal formula
(funicle of two distinct segments, club of
three or four segments), fore wing venation
and setation, non-spinose fore tibia, and
the elongate intervolsellar process in males
also help separate this genus. An inter-
volsellar process is absent in Pterandrophy-
salis and Mirufens.

Ceratogramma was treated as a junior
synonym of the Old World Szelenyia by
Fursov (1995a). It is resurrected here and
includes eight New World species, and the
recently described Spanish species, Cerato-
gramma tatianae (Fursov), new combina-
tion. The generic synonymy proposed by
Fursov was prompted by the discovery of
tatianae. The fact that certain features also
are shared with Szelenyia tamaricis Nowicki
prompted synonymy. Included are similar
fore wing venation and setation, similar
antennal segmentation, and the two-seg-
mented maxillary palp.

As indicated by Pinto and Viggiani
(1991) Ceratogramma is indeed similar to
Szelenyia. Synonymy may eventually be
appropriate, but at the present time I am
unable to identify clearly derived features
shared by S. tamaricis and Ceratogramma
species, nor did Fursov (1995a) propose
any. Whereas Ceratogramma, as defined
here, is convincingly monophyletic, the
concept of Szelenyia offered by Fursov
renders the genus difficult to define except
by a combination of generalized and likely
primitive features. Thus, the two defining
traits of Ceratogramma, the three anellar
segments and a prolonged intervolsellar
process (male unknown in C. tatianae,
however), are unique synapomorphies not
shared by S. tamaricis. Features used to
argue a tie to S. tamaricis, also are charac-
teristic of other primitive trichogrammatid
genera. The limited material of Szelenyia

available for this study suggests that there
are undescribed species in Europe and
Africa similar to S. tamaricis. Also, as
indicated below, Szelenyia is considerably
more similar to Pterandrophysalis than to
Ceratogramma. Pending a study of this
entire generic complex it seems more
practical to retain original definitions
which at least allow the unambiguous
identification of both Szelenyia and Cerato-
gramma (see Doutt and Viggiani 1968, Pinto
and Viggiani 1991).

New World records. — Argentina: Neu-
quen. Brazil: Santa Catarina. Chile: Re-
gions VIII, IX, X. Colombia. Costa Rica.
Ecuador. French Guiana. Guatemala. Pan-
ama. Peru. United States: Florida (intro-
duced), New Mexico, Texas, Utah. Vene-
zuela. West Indies.

Hosts. — Coleoptera: Curculionidae (Del-
vare 1988).

Ceratogramma jeffersi Pinto, new species

(Figs 13, 14, 100, 202)

Diagnosis. — Antennal shape and segment
number similar in both sexes; scape not
inflated. Fore wing without fumation; MV
shorter than both SV and PM; postmargi-
nal vein, radial process and basal vein
track absent. Mesophragma entire apically,
not notched.

Description. — Body length c. 1.0 mm.
Color uniformly dark brown except tarsi,
base and apex of tibiae and antennal
flagellum considerably lighter, also with
a slightly lighter midline on mesoscutum
and scutellum. Fore wing disk clear,
without fumation. Female. Head about
a third wider than long, slightly wider
than mesosoma. Toruli placed at middle of
upper face. Mandible with 2 strong poste-
rior teeth, ental surface crenulate anterior-
ly. Maxillary palpal segments elongate,
cylindrical, 0.7 X length of mandible, seg-
ment II c. 1.5X length of I; terminal seta on
II slightly longer than length of segment.
Measurements (length/width; holotype) of
antennal scape, pedicel, Fl, F2 and club as

Volume 15, Number 1, 2006

follows: 64/23, 35/27, 31/33, 35/30, 95/35;
club symmetrical, widest near junction of
CI and C2 then tapering evenly to apex, CI
c. 0.7X the length of C2, C2 slightly shorter
than C3, C3 subcorneal and without a patch
of RS apicoventrally; PLS on all postanellar
segments; F2 and club segments each with
a single whorl of FS; Fl with one elongate
APB and a few short APA; BPS at apex of
funicular and club segments very small,
narrow, only slightly expanded apically;
longest FS c. 1.4X maximum club width;
scape not inflated. Midlobe of mesoscutum
usually with 3 pair of elongate, stout setae
(only 2 pair in female from New Mexico,
and 4 setae on left side of holotype);
scutellum with 2 pair of similar setae.
Scutellum relatively long, c. 0.7 X length
of mesoscutum. Sculpturing on mesoscu-
tum and scutellum primarily lineate with
very elongate, narrow cells, cells wider
laterally and basally on both structures.
Mesophragma entire, not notched apically.
Fore wing 1.6X as long as wide, venation
attaining 0.5 X wing length, fringe setae c.
0.1 X FWW; MV curving posteriorly at base
to meet PM, rather abruptly angling
apically to form SV; MV relatively short,
c. 0.8X length of SV and PM, resp., with 3
elongate setae dorsally; PM slightly dis-
junct from MV, with 2 strong setae, lacking
a radial process; postmarginal vein absent;
costal cell well developed, with a row of c.
10 dorsal setae along anterior margin and
several scattered ventral setae behind; disk
densely setose, setal tracks R, RS2, r-m, M
and Cul present, all except R somewhat
obscured by surrounding setae, RSI setae
not distinct from surrounding setation,
a basal vein track absent. Hind wing broad,
cultriform, posterior fringe only c. 0.6 X
HWW; disk densely setose, without dis-
tinct linear tracks. First tarsomere of mid-
leg similar in length to second and third.
Ovipositor elongate but not extending
appreciably behind apex of metanotum,
gonoplac short, OL/HTL = 1.9; hypopy-
gium extending only c. 0.5 length of
ovipositor.

Male. As female except funicular seg-
ments slightly longer and narrower, and
with FS on Fl; FS longer on all segments,
longest c. 2.7X maximum antennal width.
Genitalia 3.4 X as long as wide, VS and IVP
extending well beyond apex of PAR;
without ventral protuberances at base of
PAR. AL slightly longer than GL, AL/HTL
= 0.7.

Types. — Holotype 9 and allotype 6*-
UNITED STATES. Utah: Washington Co.,
Snow Canyon State Park, c. 1.0 mi. N.
Ivins; iii-29-1989; sweeping Rhus, Quercus,
Lycium, and Fraxinus; J. D. Pinto, collr.; in
NMNH.

Etymology. — In memory of the American
poet, Robinson Jeffers.

Material examined. — 39, 36" (all on slides).
UNITED STATES. New Mexico: Hidalgo Co.,
Animas Mts. (east slope) (Gray Ranch: Indian
Creek wash, N. of Culberson Camp), 31
25.306'N, 108° 40.525'W; viii-5-2002; SP; 1& J.
George /M. Gates. Hidalgo Co., San Luis Pass
(Gray Ranch: N. side of Pleistocene lake bed),
31 25.556'N, 108 50.057'W; SP; 19; M. Gates.
Texas: Brewster Co., Big Bend National Park,
Rosillos Mts. (Buttrill Spring); iii-17/21-1992; 19;
J. Woolley/R. Wharton. Brewster Co., Big Bend
National Park; 29° 34' 22" N, 103 16' 19" W; viii-
18/21-1992; 1J; J. Woolley/R. Wharton. Utah:
Washington Co.; I9, IS (see Types).

Comments. — Ceratogramma jeffersi is the
most generalized species in the genus,
lacking virtually all derived features char-
acterizing the other New World species
(i.e. all characters listed in Table 1 in Pinto
and Viggiani 1991 are absent). In the key to
species presented in 1991, C. jeffersi runs
unsatisfactorily to C. etiennei and C. brasi-
liense in couplet 5. The absence of a post-
marginal vein, radial process, basal vein
track and a fourth club segment in males
separate it from both.

3. Haeckeliania Girault
(Figs 2, 26, 27, 109, 189, 209)

Haeckeliania Girault 1912. Type species: Haeck-
eliania haeckeli Girault, by original designa-
tion.

Diagnosis. — Female. Antenna with 2 anelli
and a 5-segmented club; club widest at CI,
often terminating in a short terminal pro-
cess, with at least some PLS extending
beyond surface for c. half their length or
more. Maxillary palp 2 segmented. Fore
wing varying in width, commonly relative-
ly broad; venation usually not attaining half
FWL, MV gradually broadening to apex
and longer than wide, considerably less
than 3X as wide as PM (may be slightly
narrower than PM), PM slightly shorter to,
more commonly, slightly to considerably
longer than MV, diverging posteriorly to
form a distinct costal cell, SV well de-
veloped, attached to the posteroapical cor-
ner of MV; disk moderately densely setose
usually with distinct linear setal tracks, RSI
usually present, radial process present.
Midlobe of mesoscutum and scutellum each
with 2 pair of setae, notauli usually broadly
arcuate. Metasomal tergum I (petiolar) with
a transverse row of denticles.

Male. Antenna with all PLS attached to
surface at base only, spiniform, erect. GC
completely open dorsally; PAR and VS
present, PAR bifid apically; ADG usually
longer than GC.

Distribution. — Widespread except not re-
corded from Europe; greatest diversity
appears to be in tropical areas.

Diversity. — There are nine species of
Haeckeliania described; only two, H. sperata
Pinto, from the West Indies and H. minuta
Viggiani, from Brazil, occur in the New
World. The remainder are from Asia and
Australia. Haeckeliania sperata was recently
introduced into Florida for control of
Diapirepes root weevil (Hall et al. 2001). As
indicated by Pinto (2005), Haeckeliania is
one of the largest genera of Tricho-
grammatidae with the vast majority of
species, including many from the New
World, awaiting description.

Discussion. — Haeckeliania is distinguished
by the five-segmented club with its rela-
tively wide CI and PLS which are largely
(females) or completely (males) free of the
antennal surface, the two-segmented max-

JOURNAL OF HYMENOPTERA RESEARCH

illary palp, the transverse row of denticles
on the petiolar tergum, and fore wing
characteristics. It appears closest to Bra-
cln/ufens, which is separated by the very
short, quadrate marginal vein (as wide as
long), and the midlobe of the mesoscutum
bearing three rather than two pair of setae.
The only other New World genus with
a five segmented club and a two-segment-
ed maxillary palp is Trichogrammatella. The
latter is distinguished by its elongate
stigmal vein, spinose fore tibiae and
antennal PLS which extend minimally
beyond the surface. The Old World Ophio-
neuris and Asian Eutrichogramma are simi-
lar phenetically to Haeckeliania; they are
separable by PLS structure (all attached to
antennal surface most of their length in
both sexes) and the absence of a row of
denticles on the petiolar tergum.

As noted previously (Pinto 2005) there are
two general phenotypes of Haeckeliania spe-
cies in the New World. In one, characterized
by H. minuta, the body is compact, gibbous,
and uniformly dark brown in color. In a less
diverse group characterized by H. sperata,
the body is more slender and lighter in color.

New World records. — Argentina: Chaco,
La Rioja, Misiones, Salta. Belize. Bolivia.
Brazil: Mato Grosso, Minas Gerais, Rondo-
nia, Santa Catarina, Sao Paulo. Canada:
Manitoba, Quebec. Colombia. Costa Rica.
Ecuador. Guatemala. Mexico: Baja Califor-
nia Sur, Chiapas, Colima, Guerrero, Jalisco,
Michoacan, Nuevo Leon, Quintana Roo,
Sinaloa, Tamaulipas, Yucatan. Nicaragua.
Panama. Peru. United States: Arizona,
California, Florida, Georgia, Illinois, Mary-
land, Missouri, North Carolina, South
Carolina, Virginia, West Virginia. Vene-
zuela. West Indies.

Hosts. — Coleoptera: Curculionidae (Pinto
2005).

4. Hydrophylita Ghesquiere
(Figs 15, 114a/b, 170, 203, 204)

Hydrophylax Matheson and Crosby 1912. Type
species: Hydrophylax aquivolans Matheson and
Crosby, by original designation.

Volume 15, Number 1, 2006

Hydrophylita Ghesquiere 1946 (n. ;/. for Hi/dro-
phylax Matheson and Crosby, nee Hydrophylax
Fitzinger 1843).

Lutzimicron Costa Lima 1960. Type species:
Lutzimicron lestesi Costa Lima, by monotypy.
Doutt and Viggiani (1968, as Lutzmicron,
synonymy). Herein as valid subgenus. New
status.

Diagnosis. — Body shape slender, elon-
gate; eyes dark red in color. Female.
Antenna elongate, with 1 or 2 anelli, 2
funicular and usually 3 club segments; all
postanellar segments longer than wide;
funicular segments and often club seg-
ments loosely articulated; PLS absent from
funicle; BPS extremely elongate, narrow,
often apically attenuate; apex of female
club with two large unsocketed spatulate
sensilla. Mandible with a very large poste-
rior spine. Maxillary palp 1 or 2 segment-
ed. Midlobe of mesoscutum and scutellum
each with 2 pair of strong setae. Propo-
deum elongate, almost as long as to
distinctly longer than scutellum, subrec-
tangular. Fore wing elongate, narrow, 5-
10 X as long as wide; disk sparsely to
extremely densely setose (sparsely setose
wings with setae linearly arranged, dense-
ly setose wings without linear tracks).

Male. Antenna with funicular segments
longer; club longer with segments less
closely appressed. GC completely open
dorsally, with distinct PAR and VS, IVP
absent.

Distribution. — New World, southern
Africa, SE Asia, Australia. Previously
known only from the New World.

Diversity. — Three species, all New
World. See Discussion.

Discussion. — Hydrophylita cannot be con-
fused with any other genus. Although
superficially similar to certain Oligositini
such as Prestzuichia and Sinepalpigramma, it
is easily separated by the two-segmented
funicle (one segmented in oligositines),
very dark but reddish eyes (black in
oligositines), two pair of setae on the
mesoscutellar midlobe and scutellum (one
pair on both in oligositines) and the

generalized male genitalia (highly reduced
in oligositines). The two large sensilla at
the apex of the female club (Fig. 15) and the
greatly enlarged posterior mandibular
spine in both sexes (Fig. 170) also help
distinguish the genus.

There are three species of Hydrophylita
described, all from the New World. H.
aquivolans from the United States and
Canada, and H. bachmanni De Santis from
South America have very narrow fore
wings (c. 10 x as long as wide) which taper
to a point at their apex (Fig. 114A), a one-
segmented maxillary palp, and a propodeal
length subequal to that of the scutellum.
They also have a uniquely shaped aedea-
gus which is enlarged and spatulate
apically (see Viggiani 1971, Fig. 6). The
two are distinguished by the number of
anelli - one in H. aquivolans, two in H.
bachmanni. H. lestesi Costa Lima and related
undescribed species, also from South
America, differ considerably from both by
the broader and densely setose fore wing
(c. 5x as long as wide) which is rounded
apically (Fig. 114B), a much longer propo-
deum (> length of scutellum), exserted
and spiniform PLS in females, a two-
segmented maxillary palp, and an unmod-
ified aedeagus. H. lestesi was placed in its
own genus, Lutzimicron, by Costa Lima
(1960) but subsequently synonymized with
Hydrophylita by Doutt and Viggiani (1968).
It is herein treated as a valid subgenus
(new status) based on the considerable
morphological differences.

The nominate subgenus is known only
from the New World; all extralimital
collections of the genus represent Lutzimi-
cron. Both subgenera occur throughout
Central and South America. I am not aware
of records from Mexico, and only the
nominate subgenus is recorded north of
there in SE Canada and NE United States.
In at least one New World species of
Lutzimicron C2 and C3 are partially fused
in females; in females of undescribed
species from Madagascar and Indonesia
these segments are completely fused and

66 Journal of Hymenoptera Research

the club is two segmented. The fore wing times difficult to discern in species with

also may be wider in certain extralimital denser setation. Fore tibia with a row of

species (c. 4X as long as wide). thorn-like spines on dorsal surface.

New World records.— Subgenus indication Male. Antennal club with a short fourth

follows each record (H = nominate sub- segment, postanellar segments dominated

genus, L = Lutzimicron). Argentina: Bue- by elongate FS. GC open its entire length,

nos Aires (H). Belize (H/L). Bolivia (H/L). with VS and PAR, without an IVP.

Brazil: Amazonas (L), Goias (H/L), Rio de Distribution.— Widespread. Occurring

Janeiro (L). Canada: Ontario (H). Colom- from Alaska south to Guatemala in the

bia (H/L). Costa Rica (H/L). Ecuador (H/ New World, with a single South American

L). Guatemala (H). Panama (H/L). United record (Chile).

States: New York (H). Venezuela (L). West Diversity. — Sixteen species are placed in

Indies (L). Mirufens; none is from the New World

Hosts. — Odonata: Coenagrionidae [Ma- although several occur here. Because Mir-

theson and Crosby 1912, for H. (H.) ufens is relatively homogeneous morpho-

aquivolans], Lestidae [Costa Lima 1960, for logically, I hesitate formally describing

H. (L.) lestesi]. New World representatives without fur-
ther study of named Palearctic species.

5. Mirufens Girault Discussion. — Ufens and Ceratogramma are

(Figs 16, 17, 101, 102, 173, 178, 205, 206) mogt ^ confused with M^fms As in

Mirufens Girault 1915. Type species: Mirufens Mirufens, males of Ufens and certain species

dentipes Girault, by original designation. of Ceratogramma have an antennal club

Trachocera Blood and Kryger 1928. Type species: with a small fourth segment. Ufens is easily

Trachocera longicauda Blood and Kryger (ju- distinguished by the one-segmented max-

nior objective synonym of Asynacta longicanda illary palp. Also the male genitalia differ

Blood), by original designation. Doutt and considerably. In Mirufens the genitalia are

Viggiani 1968 (as valid subgenus). Neto and typical of the Trichogrammatini (Type 1) in

ITZul 199? (syn°nymized Wlth nominate that the aedeagus is separate from the

j x j KT x i ™- ^rv, / genital capsule, and the latter is completely

Pseudotracnocera Neto and Fintureau 1997 (as ° ,,,.,, , • ,

subgenus). Type species: Trachocera gundlachi °Pen dorsally. Although genital structure

Nowicki, by original designation. vanes greatlY in Ufms> the 8emtal capsule

is not open its entire length dorsally and

Diagnosis. — Relatively elongate. Female, the aedeagus is not distinct. Ceratogramma
Antenna with 2 anelli, 2 funicular and 3 and Mirufens have similar genitalic struc-
club segments; postanellar segments with ture but the former is characterized by an
relatively few FS; pedicel transversely elongate intervolsellar process which is
ridged, ridges crenulate; funicular seg- absent in Mirufens. Anellar structure also
ments distinctly separated from one anoth- separates Ceratogramma. In addition, both
er, Fl often somewhat longer than F2. Ceratogramma and Ufens lack the spinose
Maxillary palp 2 segmented. Midlobe of fore tibia, and the transversely ridged,
mesoscutum and scutellum each with 2 crenulate pedicel. Mirufens probably is
pair of elongate setae. Fore wing relatively most closely related to Pseudomirufens de-
broad, < 2X as long as wide; venation with scribed from China (Lou and Yuan 1998)
SV elongate, usually subequal to or longer and also occurring in Australia (Pinto,
than MV, basal SV width commonly unpubl.). The fewer club segments in the
similar to apical width of MV, PM sub- male, and the much shorter second funic-
equal in length to MV; disk usually ular segment separate that genus from
moderately densely setose with linear setal Mirufens. The New World Trichogramma-
tracks present including RSI, tracks some- tella may also be related (see below).

Volume 15, Number 1, 2006

Doutt and Viggiani (1968) divided Mir-
ufens into two subgenera, the nominate
subgenus and Trachocera. The nominate
group included species with the marginal
vein failing to contact the anterior border
of the fore wing, a more distinct separation
between the marginal and premarginal
veins, and a longer stigmal vein. Trachocera
included species with the marginal vein
coinciding with the anterior wing margin
and more elongate club and funicular
segments. Neto and Pintureau (1997),
noting several intermediate conditions,
did not recognize this division and instead
returned all species to the nominate sub-
genus except the Palearctic M. gundlachi
which they placed in its own subgenus,
Pseudotrachocera, primarily based on its
narrower fore wings.

New World records. — Canada: Alberta,
British Columbia, Manitoba, Ontario, Que-
bec. Chile: Region X (Isla Chiloe). Guate-
mala. Mexico: Guerrero, Jalisco. United
States: Alaska, Arizona, California, Colo-
rado, Maryland, Michigan, Montana,
North Carolina, Nevada, New Mexico,
Oklahoma, Oregon, South Carolina, Texas,
Utah, Virginia, Washington, West Virginia,
Wisconsin.

Hosts. — Hemiptera: Membracidae, Cica-
dellidae (Neto and Pintureau 1997).

6. Pachamama Owen and Pinto

(Figs 18, 19, 103)

Pachamama Owen and Pinto 2004. Type species:
Pachamama speciosa Owen and Pinto, by
original designation.

Diagnosis. — Female. Compact, gibbose in
shape. Antenna with 2 anelli and a 3-
segmented club; C3 much longer than CI
and C2 combined; scape elongate, c. 3X as
long as wide. Mandible with a socketed
anterior tooth. Maxillary palp 1 segmented.
Fore wing broad, c. 1.8X as long as wide,
posterior margin with a distinct preretina-
cular lobe, venation extending c. 0.7 FWL,
fringe setae c. 0.3 X FWW; venation con-
fluent, sigmoid with greatest curvature at

base (from CS to MV), SV relatively short
and diverging little from MV and wing
margin; disk sparsely setose, setae primar-
ily restricted to apical third of wing and
not clearly arranged in linear tracks poste-
rior to the r-m, RSI absent, costal cell large
but without setae. Hind wing broad,
subtriangular, anterior margin broadly V-
shaped basal to hamuli. Both fore and hind
wing fumate except at apex. Mesophragma
not notched apically. First metasomal
tergum without a transverse row of denti-
cles.

Male. Antenna with scape extremely
enlarged, suboval, c. 1.5X as long as wide,
its width greater than length of club. GC
trough-like, completely open dorsally, with
distinct PAR and VS, IVP absent.

Distribution. — Central and NW South
America.

Diversity. — Monotypic.

Discussion. — Pachamama is one of the
most easily recognizable genera of Tricho-
grammatidae. Its gibbose body shape,
three-segmented club with C3 longer than
CI and C2 combined, the sexually di-
morphic scape (enormously inflated in
the male), and fumate fore wing with
basally sigmoid venation and a distinct
preretinacular lobe will separate it from all
other genera. Pachamama is not obviously
related to any other group. Other Tricho-
grammatini genera sharing its compact
and robust body shape include Cerato-
gramma, Poropoea, and most Haeckeliauia.
Antennal structure alone separates all
three; the former two have a distinct
funicle, and all have at least five postanel-
lar segments in both sexes. Also, unlike
Pachamama, all three genera have a two-
segmented maxillary palp. Certain species
of Adryas also may be confused with P.
speciosa. In addition to lacking the strongly
dimorphic antenna and preretinacular lobe
on the fore wing, venation in Adryas
species easily separate them. In Adryas the
venation is not sigmoid basally; instead it
is quite straight with the premarginal vein
approaching the anterior wing margin (cf.

Journal of Hymenoptera Research

Figs 103, 140). Male genitalia also are
greatly reduced in Adryas (Type 2a) which
places it in the Oligositinae.

Records. — Costa Rica. Ecuador. Hon-
duras. See Owen and Pinto (2004) for
specific collection records.

Hosts. — Unknown.

7. Paratrichogramma Girault

(Figs 28, 29, 110, 188, 210, 211)

Paratrichogramma Girault 1912. Type species:
Paratrichogramma Cinderella Girault, by origi-
nal designation.

Diagnosis. — Female. Antenna with only 3
or 4 flagellomeres: 1 or 2 anelli, 1 funicular
and 1 club segment; club with a patch of RS
apicoventrally; Fl abruptly narrowed at
apex. Mandible without a socketed anterior
tooth. Fore wing slightly more than 2x as
long as wide; venation short, abbreviated,
attaining 0.4 FWL, MV placed slightly
posterior to anterior wing margin, SV
reduced to a small sessile appendix of
MV; disk sparsely to moderately densely
setose with very few setae in basal half,
linear setal tracks present apically, RSI
absent. Mesophragma entire, not notched
apically. Metasomal tergum I (petiolar)
with a transverse row of denticles.

Male. Antenna with 2 funicular segments,
F2 abruptly and asymmetrically narrowed
apically ('bottle shaped'), club without
patch of RS ventroapically. GC with a nar-
row transverse bridge dorsal to ADG at
middle, PAR and VS present, PAR bifid
apically; ADG subequal in length or only
slightly longer than GL, AAP fused basally.

Distribution. — In New World from SW
United States S to Costa Rica with a single
record from Argentina. Also known from
Australia, southern Africa, Israel and India.

Diversity. — Eight species of Paratricho-
gramma have been described. Only one, P.
californica Doutt, occurs in the New World.
All New World collections examined are
conspecific or very close to P. californica.

Discussion. — Paratrichogramma cannot be
confused with any other genus. Its anten-

nal formula (Figs 28, 29) and distinctive
fore wing (Fig. 110) allow easy recognition.
Its fore wing venation is superficially
similar to that in Aphelinoidea and Tumidi-
clava, but in these genera the marginal vein
contacts the anterior wing margin, and the
disk is more densely setose and lacks linear
setal tracks. Paratrichogramma may be clos-
est to the Australasian Brachyia and Aits-
tralufens. All three have the aedeagal
apodemes fused basally (Fig. 210) as well
as a patch of RS on the club. Brachyia has
similarly abbreviated venation.

New World records. — Argentina: For-
mosa. Costa Rica. Mexico: Baja California
Sur, Campeche, Jalisco, Michoacan, Nuevo
Leon. United States: Arizona, California,
Texas, Utah.

Hosts. — Lepidoptera: Gracillariidae (Hay-
at and Viggiani 1984), Noctuidae (Viggiani
1976a), Lycaenidae (Pinto, unpubl.).

8. Poropoea Forster

(Figs 20, 104)

Poropoea Forster 1851. Type species: Poropoea
stollwerckii Forster, by monotypy.

Poropoeoides Nowicki 1936 (as subgenus). Type
species: Poropoea bohemica Nowicki, by mono-
typy.

Diagnosis. — Large for family, commonly
1 mm or more in length, gibbose in shape.
Female. Antenna with 2 anelli, 2 funicular
and 3 club segments; funicular segments
longer than wide, loosely articulated, fu-
nicle elongate varying from almost as long
to distinctly longer than club. Maxillary
palp 2 segmented. Mesophragma relatively
short, not notched apically. Fore wing
broad, oblate apically, < 2X as long as
wide; venation narrow, veins confluent,
strongly sinuate, with SV elongate and
almost perpendicular to MV, MV not
contacting anterior margin of wing, its
width subequal to that of SV; disk sparsely
to moderately densely setose, linear setal
tracks distinct including RSI.

Male. Antenna with all postanellar seg-
ments longer than wide, usually loosely

Volume 15, Number 1, 2006

articulated and not obviously consolidated
into a distinct funicle and club; segments
without whorls of elongate FS. GC com-
pletely open dorsally, basal margin in-
distinct, with VS, PAR and an elongate
IVP; ADG somewhat longer than GC.

Distribution. — Widespread except not re-
corded from Australia. Known in the New
World from E Canada to Brazil.

Diversity. — Poropoea includes 14 described
species. Only one, P. attelaborum Girault,
from the eastern United States, is described
from the New World. All others are from
Europe and Asia. A small number of appar-
ently undescribed species occur in the New
World. See Subba Rao (1969) and Kobayashi
and Kato (2004) for keys to species.

Discussion. — Poropoea is immediately rec-
ognizable by its fore wing venation. It is
displaced from the anterior margin of the
wing, confluent and strongly sinuate with
the stigmal vein virtually perpendicular to
the marginal vein. This coupled with
antennal structure insures identification.
Mirufens and Ceratogrnmma are somewhat
similar to Poropoea. Poropoea is separated
from both by its narrow and sinuate
venation, and its unconsolidated club in
Mew World males. It is further separated
from Mirufens by its unnotched meso-
Dhragma and male genitalia with an inter-
volsellar process. From Ceratogramma it
also differs in having two, not three anelli.

Nowicki (1936) divided the genus into
subgenera, the nominate and the mono-
:ypic Poropoeoides. The latter, based on the
European P. bohemica, was distinguished
ay the consolidated club in males. As
ndicated, in most Poropoea the club and
unicular segments are loosely articulated.
Doutt and Viggiani (1968) did not recog-
aize the distinction taxonomically, but did
aot formally synonymize Poropoeoides.

New World records. — Belize. Brazil: Ama-
wnas. Canada: Quebec. Costa Rica. Ecua-
dor. United States: Florida, Missouri. West
Indies.

Hosts. — Coleoptera: Atellabidae (Girault
L911c, Nowicki 1936, Kobayashi and Kato

2004). Both subgenera are known from
atellabids.

9. Pterandropliysalis Nowicki

(Figs 22, 23, 105, 106, 207)

Pterandropliysalis Nowicki 1935. Type species:
Pterandropliysalis levantina Nowicki, by origi-
nal designation.

Diagnosis. — Female. Antenna with 2 an-
elli, 2 funicular and 3 club segments;
funicular segments distinctly separated
from one another and from club. Fl anelli-
form, much wider than long; F2 subqua-
drate, with 2 PLS, c. 4x as long as Fl, and
c. 1.3X as wide; club widest near junction
of CI and C2, both segments subequal in
length, C3 slightly shorter, subcorneal.
Maxillary palp 2 segmented, segment I
shorter than II but distinctly wider. Fore
wing slightly more than 2x as long as
wide; venation with MV short, shorter than
PM, of normal width, SV distinct, con-
stricted between stigma and MV, radial
process present; disk densely setose, with-
out distinct setal tracks, RSI absent.

Male. Antenna with Fl somewhat larger
than in female, subequal in width to F2, F2
shaped as in female but smaller and
without PLS. Fore wing with MV and PM
consolidated, enormously inflated, occupy-
ing almost entire basal width of wing. GC
completely open dorsally, with PAR and
VS, IVP absent; ADG longer than GC.

Distribution. — Europe, north Africa and
Middle East; known from central Califor-
nia (see Discussion).

Diversity. — Monotypic.

Discussion. — Pterandropliysalis is easily
separated from all other Tricho-
grammatidae. The extent of venation in-
flation in the male fore wing is unique (cf.
Figs 105, 106). Although an inflated vena-
tion is known in males of certain other
genera (e.g. Adryas), in none of these does
the expansion span almost the entire basal
width of the wing. In the New World
Pterandropliysalis is most similar to Cerato-
gramma. Males are easily separated by fore

Journal of Hymenoptera Research

wing venation. In addition, the genital
capsule of Pterandrophysalis lacks an inter-
volsellar process, a prominent feature in
Ceratogramma. Both sexes of Pterandrophy-
salis can be distinguished by the antennal
formula. Whereas Ceratogramma has three
anelli, Pterandrophysalis has only two.

Pterandrophysalis appears to represent
little more than a species of Szelenyia with
strongly modified venation in males. All
other characters suggest synonymy. The
male genitalia are nearly identical (Vig-
giani, 1971, 1984), wing venation (exclud-
ing males of P. levant ina), discal setation of
both pairs of wings, and antennal formula
also are similar in both (see Doutt and
Viggiani 1968). Synonymy is not formal-
ized here, however, pending a more com-
plete study of Old World taxa. Also see
Ceratogramma above.

Pterandrophysalis is monotypic. The type
species, P. levantina, described from Turkey
and Syria (Nowicki 1935) also is known
from Italy, Greece and Tunisia (Pinto,
unpubl.). Recent collections by S. L. Hey-
don (UCDC) from the Cold Canyon Re-
serve in Solano Co., California, can not be
separated from P. levantina. Its occurrence
in California represents a likely Old World
introduction possibly associated with the
release of various exotic curculionid spe-
cies for control of yellow starthistle (Cen-
taurea solstitialis L.). P. levantina has been
collected in Greece from eggs of Bangaster-
nus orientalis (Capiomont) (Pinto, unpubl.);
the latter is one of the Old World species of
weevil introduced and now apparently
established in California (Pitcairn 1996).

New World records.— United States: Cali-
fornia.

Hosts. — Coleoptera: Curculionidae (un-
publ.).

10. Soikiella Nowicki

(Figs 7, 8, 30, 31, 111,212-214)

Soikiella Nowicki 1933. Type species: Soikiella
mongibelli Nowicki, by original designation.
Velten and Pinto 1990 (generic review).

Diagnosis. — Female. Antenna with 1 anel-
lus, 1 funicular and 3 club segments, club
with a short terminal process; C2 and C3
asymmetrical, C3 asymmetrically nar-
rowed to apex, dorsal surface straight from
base to apex, ventral surface convex with
a patch of RS apically; postanellar seg-
ments with few PLS attached to the
antennal surface almost their entire length;
club segments with FS. Midlobe of meso-
scutum and scutellum each with two pair
of elongate setae. Fore wing broad, c. 1.6X
as long as wide; venation short, attaining
c. 0.4 FWL, PM and MV confluent, SV
short and broad with only a slight con-
striction between stigma and MV; disk
moderately densely setose with linear setal
tracks but RSI absent. Metasomal tergum I
(petiolar) with a transverse row of denti-
cles. Ovipositor short, broad, its length
<0.75 HTL.

Male. Antenna differing considerably
from female: with 2 funicular and 3 club
segments, club segments symmetrical,
C3 without a terminal process; all post-
anellar segments with several stout,
erect, spiniform PLS. GC broad, deep,
PAR and VS present; ADG appearing
distinct from GC under light microscope
but actually connected at its sides; AAP
almost perpendicular to body of ADG (see
Discussion).

Distribution. — Western North America,
Asia and Europe.

Diversity. — There are three species of
Soikiella, S. occidental is Velten and Pinto,
from the United States and Canada, S.
mongibelli from Europe and the Middle
East, and S. asiatica Lou and Yuan, from
Asia.

Discussion. — Soikiella is separated from
other New World genera by its sexually
dimorphic antennae, and fore wing struc-
ture. The single funicular segment coupled
with three club segments as found in
females occurs only in Pintoa, the Oligosi-
tini and Bloodiella. Its broad fore wing
separates it from the former two, and the
absence of an RSI separates it from the

/olume 15, Number 1, 2006

atter. In addition, these other taxa lack
iimilarly sexually dimorphic antennae.

Soikiella is not obviously related to any
)ther genus. Fore wing and female anten-
lal structure suggest affinity to Bloodiella,
lowever the male antennae and genitalia
ire considerably different in the two
Velten and Pinto 1990). The position of
he apodemes at approximate right angles
o the aedeagus (Fig. 213) is a feature
cnown only in this genus. Soikiella is
omewhat annectant between the Tricho-
;rammatinae and Oligositinae, and place-
nent in the former is arbitrary. As in-
licated earlier, the aedeagus appears dis-
inct from the genital capsule under the
ight microscope and in certain views with
-EM examination as well (Fig. 212) sug-
;esting Trichogrammatinae. Other SEM
•reparations show possibly flexible lateral
onnections to the apex of the capsule
Fig. 213) suggesting oligositine affinity.
Tiis feature does not resemble the com-
pete aedeagus/genital capsule consolida-
ion typical of the Oligositinae however.
oikiella is placed in the Trichogrammati-
tae for this reason and the fact that the
?rgum of its petiolar segment bears a trans-
erse band of denticles as do several other
enera in that subfamily. The molecular
vidence also suggests intermediacy to the
wo subfamilies (Owen et al., in prep.).

New World records. — Canada: Alberta,
British Columbia. United States: Arizona,
"alifornia, Texas, Washington, Wyoming.

Hosts. — Diptera: Asilidae (Velten and
'into 1990).

11. Thanatogrammn Pinto, new genus

(Figs 32, 33, 112, 215-217)

Type species. — Thanatogramma oweni Pin-
3, n. sp.

Description. — Female. Antenna with 2
nelli, and a 2-segmented funicle and club,
unicular segments well separated, F2
horter than Fl, club segments closely
ppressed, C2 with apicoventral surface
onvex, with a patch of RS. Mandible with

3 posterior teeth and 1 elongate, socketed
anterior tooth. Maxillary palp 1 segmented.
Midlobe of mesoscutum and scutellum each
with 2 pair of moderately long setae (broken
in female, based on males); propodeum
narrow, only slightly longer than metano-
tum, strap-like, not modified into a longer
or elevated discal area at middle; meso-
phragma not notched apically. Fore wing
relatively wide, less than 2x as long as
wide, fringe setae short, the longest c. 0.1 X
FWW; venation with MV slightly longer
than PM, PM and MV disjunct, SV well
developed with a constriction between
stigma and MV; disk moderately densely
setose, with setal tracks poorly indicated,
RSI absent, setation extending basally to
area behind MV. Hind wing with 3 linear
setal tracks. Metasomal tergum I (petiolar)
with a transverse row of denticles. Ovipos-
itor short, length less than HTL.

Male. Differing from female in antennal
structure. Club 3 segmented and funicular
segments longer and wider. Genitalia
generalized: ADG distinct, not fused to
GC and considerably longer, aedeagal
AAP not fused at base; GC with distinct
VS and PAR, IVP absent; GC not complete-
ly open dorsally, with a relatively short
subtriangular sclerotized DLA near apex
and above the ADG forming the posterior
border of an elongate ADA.

Etymology. — Death drawing or image.
From, Thanatos, ancient Greek god of
death; gramma (Gr.), common ending for
trichogrammatid genera signifying 'lines',
'that which is drawn', etc.; gender neuter.

Distribution. — Known only from a single
locality in eastcentral California.

Diversity. — Monotypic.

Discussion. — The sexually dimorphic an-
tennal formula (2F, 3C in ',_?; 2F, 2C in 9)
along with characteristics of the funicle (Fl
and F2 distinctly separated, relatively
elongate and loosely articulated), fore wing
(relatively broad [< 2x as long as wide],
short fringe setae [c. 0.1 X FWW], distinct
SV with constriction between stigma and
MV), and maxilla (single segmented) sep-

Journal of Hymenoptera Research

arate Thanatogramma from all other tricho-
grammatid genera.

The new genus is most similar to
Trichogrammatomyia, and the Australasian
Brachyia and Australufens all of which have
similar antennal structure. From Tricho-
grammatomyia it is separated by its broader
wing and much shorter fringe setae (cf.
Figs 112, 115), as well as the longer
parameres relative to the volsellae (cf.
Figs 217, 222). Also all species of Tricho-
grammatomyia except T. tortricis have only
a single club segment. These features also
separate Australufens. From Brachyia, Tha-
natogramma is separated by its one-seg-
mented maxillary palp and distinct stigmal
vein (the stigma is abbreviated and sessile
to the MV in Brachyia). The male genitalia
of Brachyia have a similarly elongate
aedeagus but its apodemes are fused
basally, not separated as in Thanatogramma.
Also, Thanatogramma is the only one of
these genera to have a socketed anterior
mandibular tooth, an apically unnotched
mesophragma and a dorsal lamina above
the aedeagus. These features also occur in
Trichogramma, a genus differing consider-
ably in fore wing structure (cf. Figs 112,
113).

Records.— United States: California (see
below for specific records).

Hosts. — Unknown.

Thanatogramma oweni Pinto, new species

Description—Body length c. 0.6 mm
(from slides), color uniformly dark brown
except tarsi, base of tibiae and vertex of
head slightly lighter. Female (N = 2). Anten-
na (Fig. 32) with scape elongate, subequal
in length to club, 4.3 X as long as wide;
pedicel half the length of scape, c. twice as
long as wide; funicular segments subqua-
drate, each segment less than half as long
as pedicel, Fl as long as wide, slightly
longer than F2, F2 wider than long, both
segments equally wide, only half the
maximum width and 0.4X maximum
length of club; A2 closely appressed to

Fl; club 2.7X as long as wide, suboval, C2
twice the length of CI, with apicoventral
patch of RS occupying 0.6 its length; funicle
and club covered with moderately dense
and stiff APA; funicle without PLS; club
with 1 and 6 PLS on CI and C2, re-
spectively; a single short subglobose BPS
at apex of Fl, F2 and CI; UPP at apex of C2
setiform, about the same length as adjacent
RS; FS absent from funicle, sparse on club.
Mandible with socketed anterior tooth less
sclerotized but as long as unsocketed
posterior teeth. Maxillary palp slightly-
over twice as long as wide, apical sensillum
spiniform, c. half the palpal length. Meso-
soma with relatively large, finely etched
reticulae dorsally, without obvious intrar-
eticular sculpturing; midlobe of mesoscu-
tum c. 1.75 X the length of scutellum;
propodeal spiracles at anterior edge of
tergum; tarsi shorter than tibiae, all apical
tibial spurs distinctly shorter than first
tarsomere. Fore wing (Fig. 112) 1.8 X as long
as wide; venation extending c. 0.5 FWL,
longest fringe setae c. 0.1 X FWW, MV c.
1.2X length of PM and c. 2.7x as long as
wide, MV slightly wider than PM, PM with
2 setae and 2 campaniform sensilla at apex,
SV slanted posteroapically with a distinct
constriction between MV and stigma, costal
cell well developed, with 2 setae. Hind wing
with 3 linear setal tracks, scattered setae
between middle and posterior tracks, pos-
terior track not attaining apex of wing.
Ovipositor short, broad, OL/HTL = 0.66;
hypopygium not apparent.

Male. Differing from female as follows:
Antenna (Fig. 33) with club 3 segmented;
funicular segments both slightly longer
than wide, subequal in length and width,
each segment c. 0.7 X length of pedicel,
funicle c. 0.7 X the length and 0.8 X the
width of club; club distinctly longer than
scape, club segments more loosely associ-
ated, CI slightly longer than wide, C2
slightly longer than other club segments, c.
1.6X as long as wide, C3 subcorneal, 1.8 X
as long as wide; elongate FS dominating
antennal surface of all postanellar seg-

'olume 15, Number 1, 2006

rtents, representing the only setiform
tructures except for a single ATB at apex
>f CI, longest FS c. 2X maximum club
vidth; 1 PLS present on each club segment.
Genitalia (Figs 215-217) with GC 2.3 X as
ong as wide, PAR extending considerably
>eyond VS, IVP absent, basal margin of GC
ndistinct, ADA elongate, c. 0.7 the length
>f GC, DLA short, c. 0.2 x GC length; ADG
.6X as long as GC, AAP comprising at
east 0.5 length of ADG, not fused basally,
VL/HTL 0.82-1.05 (N = 2).

Types.— Holqtype 9. UNITED STATES. Cali-
yrnia: Inyo Co., Panamint Valley, Warm Sulfur
iprings; 36 07.15' N, 117 12.91' W; v-17-2002;
'YPT near mesquite and Allenrolfea" '; A. Owen/
. Pinto, collrs.; in NMNH. Allotype > Same as
lolotype except - 36 07.159' N, 117 1 2.874 'W;
ii-21/22-2002; "YPT around tamarisk and mes-
|uite"; J. George/A. Owen, collrs.; also in
sIMNH. Two paratypes as follows: I9, data
ame as holotype except labeled as - Indian
lanch Road, c. 1.5 mi. S; Warm Sulphur
iprings; 36 07.48' N, 117 12.83' W; v-15-2003;
creen sweep mesquite, Allenrolfea, etc.; J. Pinto,
ollr; UCRC. 16*, locality as 9 paratype except no
oordinates indicated and collected v-20-1991;
'screen sweeping Prosopis and Anemopsis"; J.
'into, collr.; UCRC.

Etymology. — After Albert K. Owen, who
nade considerable effort to collect this
mcommon species.

Comments. — T. oweni is known only from
he type material. The female paratype is
1 molecular voucher (Owen et al., in prep.)
ind is represented on a slide by a fore wing
tnd an antenna.

12. Triclwgramma Westwood

(Figs 9, 34, 35, 113, 169, 194, 198, 199, 218,

219)

rrichogramma Westwood 1833. Type species:
Trichogramma evanescens Westwood, by
monotypy. Sorokina 1993 (key to world
species). Pinto 1999 (revision of North Amer-
ican species).

Zalleptiles Haliday 1833. Type species: Calleptiles
latipennis Haliday, by monotypy.

Pentarthron Packard 1872. Type species: Tricho-
gramma minutum Riley, by monotypy.

Aprobosca Westwood 1878 (as subgenus). Type

species: Trichogramma (Aprobosca) erosicomis

Westwood, by monotypy.
Oophthora Aurivillius 1897. Type species:

Oophthora semblidis Aurivillius, by monotypy.
Xanthoatomus Ashmead 1904. Type species:

Xanthoatomus albipes Ashmead, by monotypy.
Neotrichogramma Girault 1911. Type species:

Neotrichogramma acutiventre Girault, by orig-
inal designation.
Trichogrammatana Girault 1932 (as subgenus).

Type species: Trichogramma singularis Girault^

by monotypy.
Trichogrammanza Carver 1978 (as subgenus,

currently valid). Type species: Trichogramma

funiculatum Carver, by original designation.
Nuniella Kostadinov 1988. Type species: Nuniella

bistrae Kostadinov, by original designation.

Pintureau 1993 (as synonym).
Vanlisus Pinto 1992 (as subgenus, currently

valid). Type species: Trichogramma lachesis

Pinto, by original designation.

Diagnosis. — Female (all subgenera). An-
tenna with 2 anelli, 2 funicular and 1 club
segment; club with a patch of RS apicov-
entrally; A2 closely appressed to Fl;
funicular segments narrower than club;
PLS absent on funicle, those on club
attached to surface for almost entire length.
Mandible with a socketed anterior tooth.
Maxillary palp with 1 or rarely (Vanlisus)
with an apparently small apical second
segment. Fore wing moderately wide;
venation sinuate with MV curving apically
away from wing margin to form an
elongate SV, PM and MV confluent, PM
with 2 setae and 2 campaniform sensilla at
apex; disk with most setae arranged in
linear tracks, RSI present. Hind wing
having 2-3 setal tracks on disk, middle
track reaching apex of wing, anterior and/
or posterior tracks complete or not. Meso-
phragma not notched apically. Metasomal
tergum I (petiolar) with a transverse row of
erect, sharply pointed denticles.

Male. Trichogramma (Trichogramma): An-
tenna with only a single distinct postanel-
lar segment - an elongate and usually
slightly curved club segment with incom-
plete, obsolescent lines of internal division.

Journal of Hymenoptera Research

Al present; A2 fused to base of club. Club lamina on the male genital capsule (Pinto
with numerous exserted FS, PLS attached 1999). However, as indicated, a similar struc-
to antennal surface for most of their length, ture also occurs in males of Tluinatogramma.
never spiniform. GC with a distinct, often Trichogramma is the best known genus in
elongate, DLA of varying shape; VS and the family because of its use for inundative
PAR present, IVP usually present and well biological control of pest Lepidoptera
developed. Trichogramma (Vanlisus): An- (Smith 1996, Parra and Zucchi 2004). The
tenna with 2 funicular and 3 club seg- New World species are divided into two
ments; club compact, C3 subcorneal, point- subgenera, the nominate and Vanlisus (Pin-
ed at apex, sensilla as in nominate sub- to 1999). Their separation is provided in the
genus. GC as in nominate subgenus except key and in the diagnosis above. The third
ADG apparently bilobed in New World subgenus, Trichogrammanza, restricted to
species (Pinto 1992). Australia and New Zealand, is separated

Distribut ion.— Cosmopolitan. by male antennal formula (2F/1C).

Diversity.— With 210 described species The North American species of Tricho-
worldwide, Trichogramma is the largest gramma were recently revised by Pinto
genus of Trichogrammatidae. Of this total, (1999). The South American fauna has yet
98 are from the New World (47 Nearctic, 35 to be studied in detail. Important recent
Neotropical, 16 both regions). Three spe- treatments of South American species in-
cies belong to the uncommonly collected elude Zucchi and Monteiro (1997), and
subgenus Vanlisus. The remainder are Querino and Zucchi (2003a, b, c).
assigned to the nominate subgenus. The Nezv World records. — Because of the cos-
third subgenus, Trichogrammanza, occurs in mopolitan distribution of this genus, dis-
Australia and New Zealand. Although tribution records are not included. In
relatively well studied compared to other general, the nominate subgenus occurs
genera, undescribed species remain in all virtually everywhere. Vanlisus is known
parts of the world. from Central America, Venezuela, Brazil

There has been considerable artificial and Mexico. Details on species distribu-

movement of Trichogramma species for bi- tions can be found in Pinto (1999), and in

ological control. Although there are excep- Zucchi and Monteiro (1997).

tions (e.g. Pinto et al. 1993), there is little Hosts. — TricJiogramma species primarily

evidence that most exotic species establish are parasitoids of Lepidoptera but eggs of

in areas of introduction (Pinto 1999). several other orders, Diptera, Hymenop-

Discussion. — The characteristic sinuate fore tera (Symphyta) and Neuroptera in par-
wing venation with an RSI setal track and 2F/ ticular, are attacked as well. Pinto (1999)
1C antennal formula in females separate itemizes known hosts for the North
Trichogramma from other genera. The genus is American species, and Zucchi and Mon-
most similar to Trichogrammatoidea. Separation teiro (1997) list hosts for those in South
of the two is covered in more detail in the America,
treatment of the latter. Other New World

genera that may be confused include Para- 13. Trichogrammatella Girault

trichogramma and Trichogrammatomyia. As in (Figs 21, 107, 208)

Trichogramma females of both have a single ^ • , , „ r. ,, 1n11 T

cS & Tricho^ramniatella Girault 1911. Type species:

antennal club segment with a patch of recurv- Trichogrammatella tristis Girault, by original

ed setae ventroapically; general body shape is designation

similar as well. Both genera lack the distinctive

sinuate venation of Trichogramma. Tricho- Diagnosis. — Female. Antenna with 2 anel-

gramma has previously been characterized as lar and 5 club segments; club relatively

the only trichogrammatid genus with a dorsal narrow, widest at CI and C2; all segments

Volume 15, Number 1, 2006

with PLS; pedicel transversely ridged,
ridges usually crenulate. Maxillary palp 2
segmented. Mid lobe of mesoscutum and
scutellum each with 2 pair of elongate
setae. Fore wing moderately wide, c. twice
as long as wide; venation with PM disjunct
from MV, MV positioned slightly but
distinctly behind anterior margin of wing
and diverging apically to form an elongate
SV, SV length subequal to or greater than
MV length, not deviating strongly from
wing margin, slanted toward middle of
apical wing margin; disk moderately
densely setose with linear setal tracks
including RSI present. Fore tibia with
short, thorn-like spines on dorsal surface.

Male. Club shorter, PLS present or absent
on CI. GC apparently completely open
dorsally, with PAR and VS present, IVP
absent.

Distribution. — New World tropics.

Diversity. — Monotypic. Trichogrammatella
is uncommon in collections. Limited mate-
rial available signals at least a few unde-
scribed species.

Discussion. — This genus is easily distin-
guished by the clearly five-segmented club
in both sexes with CI and C2 subequal in
width, the transversely ridged and usually
crenulate pedicel, the placement of the
marginal vein behind the fore wing mar-
gin, and the elongate stigmal vein which
diverges modestly from the marginal vein.
Trichogrammatella is most similar to Miru-
fens. Both have similar fore wing venation,
a transversely ridged pedicel, dorsally
spinose fore tibiae, and two-segmented
maxillary palp. Antennal formula sepa-
rates them. Mirufens has a distinct funicle,
and males are further distinguished by the
four-segmented club. Although the trans-
verse ridges on the pedicel of both genera
are crenulate, they are not consistently so
in Trichogrammatella.

Records. — Bolivia. Brazil: Rondonia.
Costa Rica. Ecuador. Guatemala. Panama.
West Indies.

Hosts. Hemiptera: Membracidae (Girault
1911b).

14. Trichogrammatoidea Girault

(Figs 36, 114, 220, 221)

Trichogrammatoidea Girault 1911. Type species:
Chaetostricha nana Zehntner, by original des-
ignation. Nagaraja 1978 (partial revision).

Diagnosis. — Female. Antenna with 2 an-
elli, 2 funicular and 1 club segment; club
with a patch of RS apicoventrally; funicular
segments narrower than club; PLS absent
on funicle, those on club attached to
surface almost entire length. Mandible
with a socketed anterior tooth. Maxillary
palp 1 segmented. Midlobe of scutum and
scutellum each with 2 pair of setae, anterior
pair on scutellum much shorter than
posterior pair. Mesophragma entire at
apex, not notched. Fore wing moderately
wide, 2.0-2.5 X as long as wide, fringe setae
0.2-0.5 X FWW; venation sinuate with MV
curving apically away from wing margin
to form an elongate SV, PM and MV
confluent, PM with only 1 seta and 1
campaniform sensillum at apex; disk with
most setae arranged in linear tracks, RSI
absent. Hind wing disk with an incomplete
medial setal track only. Metasomal tergum
I (petiolar) with a transverse row of erect,
sharply pointed denticles.

Male. Antenna with 2 funicular and 3 club
segments, club segments loosely joined, C3
rounded apically; most species with numer-
ous exserted and elongate spiniform PLS on
club and funicle. GC with PAR and VS
present, IVP present or not but very short if
present, without a DLA or sclerotized
bridge of any kind dorsal to ADG.

Distribution. — Primarily distributed in
the Southern Hemisphere and Asia with
one uncommon species from Europe.

Diversity. — Twenty-five species are as-
signed to Trichogrammatoidea. The number
occurring in the New World is unclear
because of several introductions for bi-
ological control (Querino and Zucchi 2004).
Of the ten species recorded only five are
indigenous, all from South America and
the West Indies. Trichogrammatoidea bactrae
Nagaraja, was introduced recently into

southwestern North America from Austra-
lia (Hutchison et al. 1990); its establishment
remains questionable.

The indigenous New World species
include: Trichogrammatoidea annulata De
Santis (Brazil), T. bennetti Nagaraja (Brazil),
T. brasiliensis (Ashmead) (Brazil), T. hypsi-
pylae Nagaraja (Trinidad and Tobago), and
T. signiphoroides Brethes (Argentina). Those
introduced are T. armigera Manjunath, T.
bactrae, T. eldanae Viggiani, T. nana (Zehnt-
ner), and T. robusta Nagaraja (Noyes 2001).
It should be mentioned that Trichogramma-
toidca brasiliensis was described as a Tricho-
gramma and that all of the numerous papers
in the Trichogramma biological control liter-
ature using this incorrect combination are
misidentifications (Pinto 1997b).

Discussion. — Trichogrammatoidea is most
likely confused with Trichogramma. Males
can always be separated by genitalia. In
Trichogrammatoidea the genital capsule
lacks the dorsal lamina found in all
Trichogramma (Fig. 218); instead, except
perhaps for a membranous sheet, it is
completely open dorsally and trough-like
(Fig. 220). Males of virtually all species also
are separated by antennal segmentation
(two funicular and three club segments in
Trichogrammatoidea; a single compound
postanellar segment in Trichogramma). The
only exceptions to this antennal difference
are species of the uncommonly collected
Trichogramma (Vanlisus) which have the
same antennal segmentation as Trichogram-
matoidea (Pinto 1999). Females and males
potentially confused with Vanlisus are best
separated by fore and hind wing traits
(Pinto 1997b). In the fore wing of Tricho-
grammatoidea the RSI is absent and the
premarginal vein has only a single seta;
also the hind wing has only a single linear
setal track. In the Trichogramma fore wing
an RSI is present, the premarginal vein has
two setae, and the hind wing has two or
three setal tracks, although all but the
middle track may be incomplete.

The review of Trichogrammatoidea and
key to species by Nagaraja (1978) does not

Journal of Hymenoptera Research

include all of the species that were known
to occur. Querino and Zucchi (2004) re-
cently redescribed T. annulata, a relatively
common South American species; they also
presented a key to the Brazilian species.

New World records. — Argentina: Tucuman.
Brazil: Bahia, Minas Gerais, Parana, Rio
Grande do Sol, Sao Paulo. Chile: Juan Fer-
nandez Islands. Colombia. Costa Rica. Ecua-
dor. Guatemala. Venezuela. West Indies.

Hosts. — Lepidoptera: recorded from eggs
of several families (see Nagaraja 1978, De
Santis 1981).

15. Trichogrammntoim/ia Girault

(Figs 37, 38, 115, 222)

Trichogrammatomyia Girault 1916. Type species:
Trichogrammatomyia tortricis Girault, by orig-
inal designation.

Diagnosis. — Small, c. 0.3 mm in length,
body shape moderately robust. Female.
Antenna with 2 anelli, 2 funicular and 1
or (less commonly) 2 club segments; club
with a patch of recurved setae along
apicoventral surface; PLS on club only.
Mandible without a socketed tooth. Maxil-
lary palp 1 segmented. Mid lobe of scutum
and scutellum each with 2 pair of setae,
anterior pair on scutellum distinctly
shorter than posterior pair. Fore wing 2.0-
2.5 X as long as wide; fringe setae elongate,
the longest c. 0.5 X FWW; venation with MV
abruptly ending apically, stigmal vein
approximately perpendicular to MV, with
a distinct constriction between stigma and
MV, MV longer than PM, a very short
postmarginal vein often apparent; disk
moderately densely setose, linear setal
tracks sometimes obsolescent in posterior
half of wing, RSI absent. Mesophragma
notched apically. Metasomal tergum I (pet-
iolar) with a transverse row of denticles.

Male. Antenna with 2 funicular and 3
club segments; with PLS on F2 and all club
segments. GC small subquadrate, com-
pletely open dorsally, trough-like, basal
margin indistinct, with VS and PAR pres-
ent, the former extending considerably

Volume 15, Number 1, 2006

beyond PAR, a poorly developed IVP
present or not.

Distribution. — Primarily New World
from SE Canada to Argentina; also re-
corded from the Oceanic Region. See
Discussion.

Diversity. — Monotypic. At least two or
three undescribed species occur in the New
World.

Discussion. — Within the New World Tri-
chogrammatomyia is most easily confused
with Thanatogramma, Trichogramma and
Trichogrammatoidea. In all, the female an-
tenna has similar segmentation with
a patch of recurved setae apicoventrally
on the club. Males of Thanatogramma,
Trichogrammatoidea and Trichogramma (Van-
lisus) also have an antennal formula iden-
tical to males of Trichogrammatomyia. The
latter is separated by fore wing, mandible
and mesophragmal structure. In Tricho-
grammatoidea and Trichogramma the mar-
ginal vein gradually curves away from the
wing margin to form the stigmal vein
(sinuate venation) and there is no abrupt
change in width between the two veins. In
Trichogrammatomyia, the marginal vein
ends abruptly, the stigmal vein is roughly
perpendicular to it and is much narrower
basally than the marginal vein; also its fore
wing is considerably narrower and the
marginal fringe considerably longer
(Fig. 115). From TJianatogramma, which
has roughly similar venation, it is separat-
ed by its narrower fore wing, considerably
longer wing fringe and much smaller size.
The absence of an anterior socketed man-
dibular tooth, its apically notched meso-
phragma (Fig. 193) and the absence of
a dorsal lamina on the male genitalia
further distinguishes Trichogrammatomyia.

Trichograuuuatomyia is very similar to the
Australasian Australufens. Body shape, an-
tennal structure in both sexes, wing struc-
ture, and male genitalia are similar in both.
The only difference I detect is that in
Australufens the aedeagal apodemes in the
male genitalia are basally convergent and
apparently fused (as in Fig. 210), whereas

they are separated and subparallel their
entire length in Trichogrammatomyia. The
Australasian genus Pseudogrammina,
a probable synonym of Australufens, is
similar. Previous authors did not compare
these genera presumably because the fe-
male of Trichogramma tonn/ia was character-
ized as having two club segments based on
the condition found in T. tortricis, its type
species, whereas only a single club seg-
ment characterizes Australufens (Doutt and
Viggiani 1968). It now appears that T.
tortricis is an exception. Females of unde-
scribed Trichogrammatomyia either have
a single club segment or only a partially
divided club.

Trichogrammatomyia and Australufens are
similar to Thanatogramma and the Austra-
lasian Brachyia. Antennal formula is com-
parable in all three. It should be noted that,
of the four, only Australufens and Brachyia
are characterized by basal fusion of the
aedeagal apodemes. Also, at least one
undescribed species of Australufens has
a two-segmented maxillary palp, a charac-
teristic of all Brachyia. Basally fused aedea-
gal apodemes also characterize Paratricho-
gramma.

Trichogrammatomyia is uncommonly col-
lected. Except for the type series I have not
seen specimens of T. tortricis in collections.
The only other records for the species are
those listed by Doutt and Viggiani (1968)
(see below); all are from SE Canada and
NE United States. The other collections
reported below represent undescribed spe-
cies. Unlike T. tortricis the females of these
species have a single club segment and the
second funicular segment is about as long
as the first (it is considerably shorter in the
type species).

In addition to the New World, Tricho-
grammatomyia also has been collected in the
Oceanic Region. Nafus (1993) records the
genus from Guam. I have seen single males
from Bora Bora (French Polynesia) and
from the Hawaiian Islands (Molokai).
Because the aedeagal apodemes in these
males do not fuse basally they are

Journal of Hymenoptera Research

not assigned to Australufens. However, I
have not seen material studied by Nafus
and cannot verify that identification. The
record of the genus from New Zealand
(Noyes and Valentine 1989) requires cor-
roboration. The fore wing illustrated is not
characteristic of Trichogrammatomyia.

New World records. — Argentina: For-
mosa, Misiones, Salta. Brazil: Para. Ca-
nada: New Brunswick (Doutt and Viggiani
1968), Ontario. Chile: Region VII. Colom-
bia. Costa Rica. Ecuador. Mexico: Baja
California Sur, Michoacan, Chiapas. Unit-
ed States: Connecticut (Doutt and Viggiani
1968), Florida, New York (Doutt and
Viggiani 1968). Venezuela. West Indies.

Hosts. — Lepidoptera: Tortricidae (Girault
1916c).

16. Viggianiella Pinto, new genus

(Figs 39, 97, 98, 116)

Type species. — Viggianiella tropica Pinto, n.
sp.

Description. — Female. Antenna with 1
anellus, 2 funicular segments and a single
club segment. Funicular segments trans-
verse; club without a patch of RS on ventral
surface. Mandible without a socketed an-
terior tooth. Maxillary palp very short,
possibly 2 segmented. Midlobe of mesos-
cutum and scutellum each with 2 pair of
elongate setae. Propodeum distinctly lon-
ger than metanotum, posterior margin
strongly undulate, disk lobiform. Wings
unique: fore wing elliptical, relatively
narrow, widest at middle, fringe setae as
long or longer than FWW; disk glabrous;
venation entirely confluent, strongly sinu-
ate, SV extremely elongate, digitiform,
unique in being directed anteroapically
and extending apically considerably be-
yond apex of MV. Hind wing narrow, disk
glabrous, with anterior and posterior fringe
setae both extremely long. Mesophragma
notched apically. Metasomal tergum I
sclerotized, without a transverse row of
denticles, anterior margin undulate, mir-
roring posterior margin of propodeum,
together delimiting a 'dumbbell-shaped'

membranous area between the two terga

(Fig. 97).

Male. — Unknown.

Etymology.— After Professor Gennaro
Viggiani (University of Naples, Portici), in
recognition of his major contributions to
our knowledge of the Trichogrammatidae;
gender feminine.

Distribution.— South America.

Diversity. — Monotypic.

Discussion. — Antennal formula coupled
with characteristics of the wings separate
this genus from all others in the family.
Fore and hind wing structure in Viggia-
niella is unique and on this basis the genus
is not easily compared to others. Fore wing
venation with its elongate, anteroapically
directed stigmal vein and the extremely
elongate anterior fringe on the hind wings
have no counterparts in the Tricho-
grammatidae.

Because males are unknown, the
placement of Viggianiella in Trichogram-
matinae is tentative and based solely
on female antennal formula and struc-
ture. Antennae characterized by two rela-
tively small funicular segments without
PLS and one club segment, occur in
females of several trichogrammatine gen-
era. Included are Apseudogramma , Austra-
lufens, Pseudogrammina, Trichogrammato-
myia, Trichogramma and Trichogrammatoi-
dea. The antenna of Viggianiella differs in
that the ventral surface of the club lacks
a patch of recurved sensilla, although what
are presumably two elongate RS occur
apically.

Records. — Brazil: Bahia. Colombia. (See
below for specific localities).
Hosts. — Unknown.

Viggianiella tropica Pinto, new species

Description. — Based on 2 slide-mounted
females. Except for body length, quantita-
tive data taken from holotype; if variable in
the two specimens data reported as a range.

Female. Body length c. 0.5 mm (N = l).
Color apparently uniformly light brown;

Volume 15, Number 1, 2006

fore wing moderately fumate except clear
at apical fifth, anterior to SV, and at base
behind SC, the latter area with a poorly
defined longitudinal dark stripe running to
base of wing. Toruli placed low on head, at
level of ventral margin of eyes. Antenna
with scape relatively narrow, slightly wid-
er in basal half; pedicel bead shaped, c. half
the length of scape; funicular segments
transverse, Fl considerably narrower and
shorter than F2; 1 -segmented club subeq-
ual in length to scape, widest at base,
gradually narrowing to apex; length/
width of scape, pedicel, funicular segments
and club: 42/12, 22/18, 6/11, 12/15, 42/16.
Antennal sensilla: anellus and Fl each with

1 APB, F2 - 2 APB, c. 3 APA and 1 PLS;
club - row of several APA at base, with
several FS (the longest c. 2.5 X greatest club
width), at least 5 PLS (3 of these extending
off surface of club for 0.3-0.5 their length),

2 very long RS? at apex extending consid-
erably beyond apex of club; BPS absent or
not visible. Maxillary palp present but very
short, possibly 2 segmented. Mandible
with 3 teeth, 2 of these strong, distinct.
Pronotum relatively long, narrowly divid-
ed longitudinally at midline. Midlobe of
mesoscutum with both pair of setae in
basal half, the two pair of scutellar setae
adjacent to posterior border; surface of
midlobe and scutellum very smooth, rela-
tively large obsolescent reticulae visible on
midlobe only; a median sulcus on scutel-
lum and basally on midlobe. Meso-
phragma elongate, c. 1.75X HTL. Fore
wing 3.8X as long as wide, widest near
middle of wing, venation attaining 0.85
FWL, fringe length 1.3-1.4X FWW; vena-
tion narrow, strongly sinuate, confluent;
relative length of PM, MV and SV =
3:8:5; MV curving basally to meet PM,
widest at apex, with 3 elongate anterior
setae; PM with 2 setae and 2 campaniform
sensilla at apex; SV without a distinct
stigma, narrowest at basal half, only
slightly widening in apical half; disk
glabrous except 1 short seta behind SC
and anterior to stigma (these setae seen

only in the female from Colombia). Hind
wing with elongate anterior fringe setae c.
0.8 X the length of posterior fringe setae,
their length 0.7 and 0.9 the distance from
hamuli to wing apex, respectively (all
measurements possible only in Colombian
specimen). Midfemur with apical spine
elongate, robust, c. half length of femur;
spines at apex of fore and hind femora
considerably shorter; tarsomeres of midleg
elongate, with segment I distinctly longer
than II or III; tarsomeres of fore and hind
legs shorter, subequal. Ovipositor not ex-
tending beyond apex of metasoma, length
2.0 X HTL, with gonangulum posterior to
base of ovipositor itself, gonoplac very
short, outer plate gradually but distinctly
narrowing to apex.

Male. — Unknown.

Type.— Holotype 9, BRAZIL. Bahia: Mu-
curi; xi-78; F. Benton; in BMNH.

Distribution. — Known only from eastern
Brazil and Colombia.

Material examined. — 2 9. BRAZIL. Bahia; lQ
(see Type). COLOMBIA. Amazonas: PNN Ama-
cayacu, Matamata (150 m); 03 23' S, 70 06' W;
Malaise trap (M682); 19; A. Parente; UCRC.

Comments. — The two females available
are not ideal for observing all important
features. For example, the maxillary palp,
visible only in the holotype, is very short
and it is not clear if one or two segments
are present. What appears to be a small
second segment may actually be the base of
the apical sensillum. Similarly, basiconic
peg sensilla (BPS) are not clearly visible on
the antenna; yet these sensilla occur in all
other trichogrammatids. Both specimens
studied show what may be a single very
elongate BPS at the middle of the club but
this must be confirmed.

17. Xenufens Girault

(Figs 40-12, 117, 223)

Xenufens Girault 1916. Type species: Xenufens
raskini Girault, by original designation.

Pseudoxenufens Yoshimoto 1976. Type species:
Pseudoxenufens forsythi Yoshimoto, by original
designation. New synonymy.

Journal of Hymenoptera Research

Diagnosis.— Female. Antenna short, com- unlike Xenufens the club usually is one

pact, "with 1 anellus, 2 funicular and 2 segmented in these genera and bears

complete club segments; terminal club a patch of RS ventrally. Trichogramma and

segment partially divided or not, without Trichogrammatoidea also have sinuate fore

a patch of RS; funicular segments trans- wing venation although the MV coincides

verse, distinct or partially fused; surface with the wing's anterior margin.

dominated by APA, without FS. Maxillary Pseudoxenufens is treated as a synonym of

palp 1 segmented. Mandible with a large Xenufens. Yoshimoto (1976) separated the

socketed anterior tooth. Midlobe of meso- two primarily by the number of funicular

scutum and scutellum each with 2 pair of segments in the female and apparently by

strong setae. Fore wing wide, c. 2X as long the placement of the marginal vein slightly

as wide; venation somewhat sinuate with behind the anterior margin of the fore

MV placed slightly behind anterior margin wing. The two are similar in both respects,

of wing, curved and narrowing apically to however. The marginal vein is similarly

form SV, SV with a distinct stigma apically; positioned, although perhaps it is slightly

disk moderately densely setose with linear more posterior in the type species of

setal tracks present including RSI. Meso- Pseudoxenufens. SEM examination of the

phragma not notched apically. Metasomal funicle of the type species of both show

tergum I (petiolar) with a transverse row of that it is only the relative length of the

denticles. funicular segments that differ. In X. ruskini,

Male. Antenna with funicular segments Fl is longer than F2 and both segments are

slightly more elongate than in female, not distinct (Fig. 40). In X. forsythi, Fl is

partially fused; FS present on both funicu- extremely short, disciform, partially fused

lar and club segments; club completely 2 to F2 and visible only in medial view

segmented (in ruskini). GC completely (Figs 41, 42). One can question whether

open dorsally, trough-like, divided apically what is being called Fl represents a funic-

into 2 lobes (fused PAR and VS?). ular or an anellar segment since neither Fl

Distribution. — Known almost exclusively nor F2 bear BPS, a feature often character-

from the New World, from SE United izing funicular segments in this family.

States S to Argentina, with single records However, the coeloconic sensillum gener-

from Papua New Guinea and Indonesia. ally associated with A2 does not occur on

Diversity. — There are two species of the putative Fl of X. forsythi (Fig. 42),

Xenufens. X. ruskini is widespread, occur- suggesting that it is a true funicular

ring throughout the range of the genus; X. segment, homologous to Fl in X. ruskini.

forsythi, new combination (from Pseudox- Also, BPS are lacking on the funicular

enufens) is known from Ecuador, Colombia segments of both species. In addition,

and Costa Rica. Single females from New structural similarity in the two species

Guinea and Indonesia in UCRC are in- suggests homology. In both, F2 is ringed

distinguishable from X. ruskini. with APA positioned slightly behind the

Discussion. — The two-segmented, highly apex of the segment, and Fl appears to lack

transverse funicular segments and two/ marginal setae or sensilla of any kind (cf.

three-segmented club, the nearly sinuate Figs 40, 42). It is possible that A2 has

fore wing venation, and placement of the merged with Fl in both species. Yoshimoto

marginal vein slightly behind the anterior also notes that the number of club seg-

wing margin separate Xenufens from all ments differs in females of Psuedoxenu-

other trichogrammatids. The genus is most fens. The difference is minimal (two in

similar to Trichogramma, Trichogrammatoi- forsythi, two with an incomplete third in

dea and Trichogrammatomyia. Females of all ruskini); overall club structure is similar

three have similar antennal structure, but in both.

Volume 15, Number 1, 2006

Males of Xenufens ruskini are virtually
unknown. Of the 160 specimens of this
species in the UCRC only one male is
represented. Males may be more common
in X. forsythi; its type series consisted of
five males and 46 females (Yoshimoto
1976). The discovery of the single male of
X. ruskini provides additional evidence for
the proposed generic synonymy. In both
species the genital capsule is divided
apically into two lobes, a unique feature
in the family [cf. Fig. 223, and Fig. Ill (4) in
Viggiani 1984].

New World records. — Argentina: Mis-
iones. Belize. Brazil: Minas Gerais, Rondo-
nia. Colombia. Costa Rica. Ecuador. Gua-
temala. Mexico: Jalisco, Tamaulipas, Ver-
acruz. Panama. Peru. United States: Flor-
ida. West Indies.

Hosts. — Lepidoptera: Nymphalidae
(Malo 1961, Yoshimoto 1976), Hesperiidae
(Girault 1916a; Pinto, unpubl.). Xenufens
has been reported as phoretic on adults of
its host. X. forsythi was taken from eggs
and adults of Caligo eurilocJius (Cramer)
(Malo 1961) and I have examined a series
of X. ruskini collected in Jalisco by A.
Warren labeled 'ex. thorax of Hesperiidae
butterflies'.

Oligositinae

Diagnosis. — Maxillary palp 1 segmented.
Male. Type 2 genitalia: GC and ADG
consolidated into a single structure and
not distinguishable, at least not in apical
half of genitalia; ADG presumably not
capable of movement independently of
GC, AAP of ADG present or not in basal
half; GC with VS and PAR present or
absent. Figs 224-259.

Discussion. — As indicated above, the defi-
nition of subfamilies adopted here transfers to
the Oligositinae the Paracentrobiini as well as
certain genera previously placed in the
Trichogrammatini. This hypothesizes a mono-
phyletic Oligositinae based on the consolida-
tion of aedeagus and genital capsule.

Paracentrobiini Howard

Diagnosis. — Antenna (Figs 43-45) with 2
funicle and 2 or 3 club segments; F2 much
shorter than Fl; PLS absent from funicular
segments in New World genera. Eyes
reddish. Midlobe of mesoscutum and
scutellum each with two pair of setae.
Male. Last visible sternum modified into
a shelf-like structure, or with a pair of
platiform or narrow and elongate appen-
dages (Figs 224-226). Type 2A genitalia:
GC subconical, bearing ventral setae, ADA
short, transverse, open dorsally; aedeagal
AAP present (Figs 224-226).

Discussion. — The Paracentrobiini include
four genera, Paracentrobia, Ittys, Ittysella,
and Paraittys. All but Paraittys occur in the
New World. Antennal structure and the
metasomal sternal modifications in males
separate paracentrobiines from all other
New World trichogrammatids. These fea-
tures are derived and the tribe is hypoth-
esized as monophyletic. The treatment of
genera below borrows from a thesis by J.
George at UCRC (George 2003).

18. Ittys Girault

(Figs 44, 119, 225)

Ittys Girault 1911. Type species: Trichogramma
ceresarum Ashmead, by original designation.
Doutt and Viggiani 1968 (as subgenus of
Paracentrobia). Viggiani 1971 (renewed gener-
ic status).

Diagnosis. — Female. Antenna with 2 an-
elli, 2 funicular and 3 club segments; Fl
longer than wide, usually cylindrical, F2
about as wide as long, not transverse; club
segments longer than wide, distinct, never
partially fused. Fore wing c. twice as long
as wide, fringe setae short, c. 0.2 x FWW;
venation with SV relatively long, with
a distinct constriction between stigma and
MV; disk moderately densely setose with
distinct linear setal tracks, RSI present,
consisting of several setae.

Male. GC with VS sickle shaped at apex;
length of ADA less than 1/3 GL; last

Journal of Hymenoptera Research

visible sternum with subcircular or sub-
triangular, platiform appendages, each
bearing a terminal spine.

Distribution. — Recorded from all regions
of the world.

Diversity. — There currently are six
named species of Ittys. Three occur in the
New World: I. ceresarum (Ashmead), I.
nympha (Girault), and /. prima Perkins.
The North American species are undergo-
ing revision by J. George (UCRC) and
current species concepts will be revised
considerably.

Discussion. — Ittys is separated from the
other New World paracentrobiine genera
by the cylindrical, longer than wide Fl and
non-transverse F2, the RSI setal track in the
fore wing, and the paired, relatively wide,
platiform appendages on the last metaso-
mal sternum in males.

New World records. — Brazil: Minas Gerais.
Costa Rica. Guatemala. Mexico: Campeche,
Chiapas, Michoacan, Nuevo Leon, San Luis
Potosi, Sinaloa, Tamaulipas, Veracruz. Yu-
catan. United States: Arizona, California,
Colorado, Florida, Georgia, New Mexico,
Texas, Indiana, Illinois, Kansas, Oregon,
Mississippi, Missouri, South Carolina, Ten-
nessee, Utah, Virginia, Washington, Wiscon-
sin, West Virginia. West Indies.

Hosts. — Hemiptera: Auchenorrhyncha
and Heteroptera (George 2003, Tipping et
al. 2005).

19. Ittysella Pinto and Viggiani

(Figs 45, 120, 226)

Ittysella Pinto and Viggiani 1987. Type species:
Ittysella lagunera Pinto and Viggiani, by
original designation.

Diagnosis. — Female. Antenna with 1 anel-
lus, 2 funicular and 2 complete club
segments, if club apparently with 3 then
CI and C2 partially fused, Fl wider than
long, F2 short, transverse. Fore wing
relatively narrow, 2.5 X as long as wide,
fringe setae elongate, c. 0.5 X FWW; vena-
tion with SV short and broad, with only
a slight constriction at base of stigma; disk

sparsely setose, with few linear setal tracks,
RSI absent or consisting of 1-3 staggered
setae.

Male. GC with VS sickle shaped at apex,
ADA length a third or more GL; last
metasomal sternal modification consisting
of a single rectangular, shelf-like structure
(not paired as in other paracentrobiine
genera).

Distribution. — Southwestern North Ame-
rica.

Diversity. — Only the type species, /. lagu-
nera, currently is assigned to Ittysella. How-
ever, additional species are known and will
soon be described by J. George (UCRC).

Discussion. — Ittysella differs from other
paracentrobiines by the single anellar
segment, and the partial or complete fusion
of CI and C2 (cf. Figs 43-45). It is further
separated from Ittys by the narrower fore
wing and its longer fringe, the poorly
developed or absent RSI, the wider than
long Fl and F2, and the undivided, shelf-
like modification of the last sternum in
males. Fore wing and funicular structure is
more similar in Paracentrobia. Unlike Itty-
sella, however, in Paracentrobia the antennal
club is three segmented, the male genitalia
lack distinct volsellae, and the modifica-
tions of the last metasomal sternum are
paired and elongate (cf. Figs 224, 226).

Neiv World records. — Mexico: Baja Cali-
fornia Sur, Coahuila, Sonora. United
States: Arizona, California, New Mexico,
Oregon, Texas, Utah.

Hosts. — Hemiptera: Cicadellidae (Pinto
and Viggiani 1987).

20. Paracentrobia Howard

(Figs 43, 118, 224)

Paracentrobia Howard 1897. Type species: Para-
centrobia punctata Howard, by monotypy.

Abbella Girault 1911. Type species: Abbella
subflava Girault, by original designation.
Doutt and Viggiani 1968 (synonymy).

Brachistella Girault 1911. Type species: Tricho-
gramma acuminatum Ashmead, by original
designation. Doutt and Viggiani 1968 (syn-
onymy, treatment as subgenus).

Volume 15, Number 1, 2006

Jassidophthora Perkins 1912. Type species: Jnssi-
dophthora prima Perkins, by monotypy. Doutt
and Viggiani 1968 (synonymy).

Abbellisca Ghesquiere 1946 (unjustified n. n. for
Abbella).

Diagnosis. — Female. Antenna with 2 an-
elli, 2 funicular and 3 club segments,
Cland C2 not partially fused; funicular
and club segments wider than long. Fore
wing relatively narrow, 2.3-2.6 X as long as
wide, fringe setae c. 0.2 X FWW; disk
moderately densely setose, with or without
distinct setal tracks, RSI usually absent.

Male. GC without distinct, sickle-shaped
VS; paired appendages of last metasomal
sternum elongate, strap shaped.

Distribution. — Worldwide.

Diversity. — Forty-three species are as-
signed to Paracentrobia, although several
of these may not be correctly placed. Eight
nominal species occur in the New World
(George 2003). The genus occurs through-
out the area but none have been described
from South America.

Discussion. — Paracentrobia is separated
from both Ittysella and Ittys by male features.
The genitalia of the latter two possess sickle
shaped volsellae (Figs 225, 226). In Paracen-
trobia volsellae are present but vestigial and
consist of obsolescent lobes on the ventral
surface; they can be seen under the SEM but
not easily with light microscopy (Fig. 224).
Male Paracentrobia also are identifiable by
the elongate, strap-like appendages on the
last metasomal sternum. Both sexes are
separated by antennal structure. Paracentro-
bia, unlike Ittysella, has two anelli and three
complete club segments. The wider than
long funicular segments provides separation
from Ittys (cf. Figs 43, 44).

Doutt and Viggiani (1968) originally
treated Ittys and Brachystella as subgenera
of Paracentrobia. Viggiani (1971) later re-
turned Ittys to generic status. Although
Brachystella remains a valid subgenus, it
never has been clearly distinguished from
the nominate group and its status is
questionable. In general, species assigned

to Brachystella have a fore wing which is
narrower and with more dense setae which
are not distinctly arranged into linear
tracks.

New World records. — Argentina: For-
mosa, Jujuy, La Rioja, Misiones, Salta.
Bermuda. Belize. Bolivia. Brazil: Goias,
Minas Gerais, Sao Paulo. Canada: Alberta,
British Columbia, Manitoba, New Brus-
wick, Ontario, Prince Edward Island. Costa
Rica. Colombia. Ecuador. Honduras. Unit-
ed States: Arizona, California, Colorado,
Georgia, Florida, Illinois, Indiana, Iowa,
Kansas, Maryland, Mississippi, Missouri,
Montana, Nebraska, New Mexico, Nevada,
North Carolina, Oklahoma, Oregon, Penn-
sylvania, South Carolina, Texas, Utah,
Virginia, Washington, Wisconsin, Wyom-
ing. Peru. Venezuela. West Indies.

Hosts. — Hemiptera: Auchenorrhyncha
(primarily) and Heteroptera (George
2003). Also recorded from Lepidoptera
(Polaszek et al. 1998) but this needs
confirmation.

Chaetostrichini Walker

Diagnosis. — Antennal segmentation vari-
able but funicle, if present and 2 segment-
ed, with F2 subequal to or longer than Fl;
PLS present or not on funicle. Eyes
reddish. Scutellum always with at least 2
pair of setae; mid lobe of mesoscutum
usually with 2 pair. Male. Last visible
metasomal sternum not modified as in
paracentrobiines (Figs 260, 261). Type 2A
genitalia: GC with or without ventral setae,
ADA relatively large and open dorsally,
aedeagal AAP present or not (Figs 227-257).

Discussion. — The definition of the Chae-
tostrichini adopted here represents all
Oligositinae without the synapomorphies
of the Paracentrobiini and Oligositini. As
such it clearly is not monophyletic. Tribal
limits are similar to those originally pro-
posed by Viggiani (1971). The only excep-
tion is the inclusion of certain genera
with Type 2A genitalia that Viggiani
originally placed in the Trichogrammatini

Journal of Hymenoptera Research

because of the presence of aedeagal apo-
demes.

Twenty-seven New World genera are
placed here. At least two assemblages
within the tribe are hypothesized as mono-
phyletic. One includes Burksiella, Centro-
biopsis, Chaetostricha, Lathromeroidea, Usca-
noidea, Zaga, and Zagella ( = Chaetostricha
group). This group is characterized as
follows: first postanellar segment (Fl or
CI) very short, transverse, considerably
shorter than and closely appressed to
second (F2 or C2); club three (rarely two)
segmented; fore wing with distinct setal
tracks including RSI; stigmal vein well
developed. Molecular evidence also sup-
ports monophyly (Owen et al., in press).
The limits of these genera are not clear-cut
and several species cannot be placed easily.
The assemblage is extremely speciose in
the New World, especially in the tropics,
with the vast majority of species unde-
scribed. Intermediate forms suggest that
current generic definitions will require
revision once species-level taxonomy is
better understood.

Another generic grouping includes Bra-
chygrammatella, Chaetogramma, and Xipho-
gramma ( = Brachygrammatella group). Char-
acteristics include the following: antennal
funicle partially or completely divided into
two subequal segments; club one or obso-
lescently two segmented and with numer-
ous short, fine APB sensilla; fore wing with
stigmal vein short and lacking a basal
constriction, disk densely setose with linear
setal tracks absent or few in number (RSI
absent). Generic limits in this assemblage
also remain confused. This problem, dis-
cussed by Pinto (1990b), is summarized
here in the treatment of Chaetogramma.

The affinity of the other genera in this
tribe are not clear. Phenetic similarities
which pose problems for identification and
certain suggested relationships are dis-
cussed in the treatments of individual
genera. The molecular study by Owen et
al. (in prep.) also considers their relation-
ships in greater detail.

21. Adelogramma Pinto, new genus

(Figs 63-66, 139, 241)

Type species. — Adelogramma primum Pin-
to, n. sp.

Description. — Body small (< 0.5 mm in
length), compact, somewhat gibbose; eyes
very dark red, approaching black in color.
Antenna strongly sexually dimorphic. Fe-
male. Antenna long compared to body
length (c. half body length in most species),
with 1 anellar segment; funicle absent; club
asymmetrically tapering from base to apex,
ventral surface broadly convex, dorsal
surface straight or slightly concave, club
segmentation obsolescent with 3 complete-
ly separated segments (C1-C3), separation
of C3 and C4 complete only over half
antennal circumference, separation of C4
and C5 even less complete, all sutures
faint; CI extremely short, anelliform; C2
transverse, much wider than long; partial
separation of C3 and C4 near base of
combined segment; club with PLS on C2
free for c. half its length and, in most
species, with at least one spiniform PLS
(entirely free of segment except at base)
and a large patch of elongate RS ventroa-
pically on C3/4/5. Maxillary palp 1 seg-
mented, with a setiform rather than a peg-
like sensillum near apex. Midlobe of
mesoscutum and scutellum each with 2
pair of elongate setae. Fore wing c. 2x as
long as wide, fringe setae elongate, 1/3 or
more FWW; disk moderately densely se-
tose with distinct setal tracks, RSI present,
with a small patch of foliate sensilla on
ventral surface anterior to retinaculum;
venation with PM disjunct and narrower
than MV, MV considerably longer, SV
distinct. Hind wing with anterior fringe
confined to apical third or fourth of wing.
Ovipositor short, length not exceeding
HTL and not extending beyond apex of
metasoma.

Male. Antenna distinctive: club suboval
with 5 segments; CI very short as in
female, C4 incompletely sclerotized on all
surfaces; C3, C4, C5 each with extremely

Volume 15, Number 1, 2006

elongate, thick spiniform PLS, their length
exceeding length of club itself. Genitalia
simple, tubular, with an elongte ADA;
AAP, PAR,VS and ventral setae absent.

Etymology. — From 'adelos' (Gr.), some-
thing imperceptible or secret; 'gramma'
(Gr.), common ending for trichogrammatid
genera signifying 'lines' 'that which is
drawn', etc.; gender neuter.

Distribution. — Eastern United States,
Central and South America.

Diversity. — There are at least four species
of Adelogramma. One occurs in the United
States; the others are distributed in Central
and South America. Only the U.S. species,
A. primum, is described below.

Discussion. — Antennal structure sepa-
rates Adelogramma from other genera. As
in certain genera of the Chaetostricha group
the antenna lacks a funicle and the club has
a very short, anelliform first segment.
However, unlike these genera, females of
Adelogramma have an asymmetrical club
with only three complete segments; also
the club usually has at least one spiniform
PLS and another which extends beyond the
antennal surface for about half its length.
The antennal club in males is unique
among New World genera. Although five
segmented, it bears several spiniform PLS
which are much longer than the club itself.
The antenna of the male of the monotypic
New Zealand genus Zelogramma described
by Noyes and Valentine (1989) bears re-
markable resemblance to that of Adelo-
gramma. Genitalic structure, however,
clearly places Zelogramma in the Tricho-
grammatinae.

Adelogramma perhaps is most similar to
Adryas. In both the antennal club is incom-
pletely five segmented in females but with five
distinct segments in males. However, in Adryas
the PLS are never spiniform in males or females,
the PM is distinctly wider than the MV and
confluent with that vein, and the hind wing
anterior fringe extends most of the distance
beyond the hamuli. Eye color also differs (bright
red in Adryas, very dark red in Adelogramma).

Records. — Colombia. Costa Rica. Ecua-
dor. United States: (see description of A.
primum for detailed records).

Hosts. — Unknown.

Adelogramma primum Pinto, new species

Description. — Female. Small, body length
0.3 mm; color entirely light brown with
antennae and legs somewhat lighter (based
on two card-mounted specimens). Fore
wing lightly fumate behind venation. Man-
dible with 3 strong, sharply pointed teeth
posteriorly and 1 shorter, less sclerotized
anterior tooth. Maxillary palp c. 3x as long
as wide, narrowing asymmetrically to
apex; terminal seta very long, c. 1.8X
length of palp. Length/width of scape,
pedicel and club: 36/10, 18/13, 50/20.
Club with a short terminal process, widest
near apex of C2, with 3 complete segments:
CI very short, anelliform, C2 transverse, c.
0.4 X as long as wide; sutures between C3
and C4 apparent on medial surface only,
a faint indication of a less complete C4-C5
suture also visible medially, all sutures
obsolescent; consolidated C3/4/5 segment
5X length of C2. Club sensilla: CI with 2
BPS; C2 with several APA, 1 elongate APB
and 1 curved PLS extending beyond surface
for c. half its length; C3/4/5 with several
APA ventrally in basal half, BPS present
adjacent to partial segment sutures and at
apex of C5, c. 7 PLS (3 of these spiniform,
attached only at base, others with apical 1/3
or so extending beyond surface), 1 APB on
dorsal surface near apex, a setiform UPP
inserted at apex of short terminal process;
all BPS large, subglobose.

Midlobe of mesoscutum and scutellum
each with 2 pair of elongate setae, anterior
pair on midlobe longer than posterior pair,
scutellar setae subequal in length; dorsum
of midlobe reticulate with irregularly
shaped (primarily subquadrate or sub-
rhomboid) cells which are widest anterior-
ly and laterally, becoming more linear and
much narrower at middle, scutellar sculp-
turing composed entirely of relatively

Journal of Hymenoptera Research

narrow, linear cells; cells on midlobe
weakly corrugated within. Propodeal disk
only slightly produced posteriorly, round-
ed, c. 2x length of metanotum. Fore wing
2X as long as wide, venation attaining 0.5
FWL, fringe setae 0.3 X FWW; venation
with MV very long, straight, 1.8X length of
PM, with 3 elongate anterior setae and 5
shorter posterior setae dorsally; PM nar-
rower, linear, angling slightly posteriorly,
with 2 elongate setae and 2 campaniform
sensilla at apex; SV well indicated, c. 1/3
the length of MV, arising at about a right
angle from posterior corner at apex of MV,
with a narrow constriction between sub-
oval stigma and MV; radial process absent.
Fore wing disk moderately densely setose
with most setae arranged in linear tracks;
RSI straight, angling posteriorly from
stigma; 2 basal track setae; narrow costal cell
with 2 setae at anterior margin; 3-4 foliate
sensilla on ventral surface anterior to reti-
naculum. Hind wing narrow, with 2 com-
plete setal tracks anteriorly and an incom-
plete posterior track (extending 0.6 distance
from hamuli to wing apex in holotype but
shorter in most other specimens). Ovipositor
short, OL/HTL = 0.80 (N=4).

Male. Antenna very different from that of
female. Club suboval, symmetrical, shorter
(only c. 0.9 length of scape), distinctly 5
segmented; CI as in female (anelliform);
C2, C3 and C4 transverse; C2 and C3
subequal in length; C4 shorter; C5 almost
twice length of C4, broadly subconical.
Club sensilla: C2-C4 each with an extreme-
ly elongate, thick spiniform PLS, C5 with 2
elongate PLS; PLS length c. 1.75 X that of
entire club; RS absent; only 1 FS visible (on
C4); 1 APB on C2 and a few APA on C2
and C3. GC short, its length c. 0.5 X HTL;
ADA occupying c. 0.5 GL.

Types.— Holotype 9 and allotype <$: UNITED
STATES. South Carolina: Dorchester Co., Harley-
ville, 10 km NE (Francis Beidler For.); bald
cypress swamp; MT/FIT; vi-1 1/23-1 987; in
CNC. One additional female designated as
paratype (data same except 'nr. Harleyville, x-
1/10-1987; MT; BRC Hym. Team'), in UCRC.

Etymology. — 'Primum' (L.), the first (Ade-
logramma to be described).

Distribution. — Southeastern United States.

Material examined. — 149, lc? (all except 29 on
slides). UNITED STATES. Florida: Fat Deer Key
(Monroe Co.); viii-31/xii-15-1986; MT/FIT; 1Q;
S./J. Peck.

Missouri: Williamsville; vii-16/viii-8-
1988, ix-1/20-1988; MT; 29; J. Becker. Louisi-
ana: Chicot State Park (Evangeline Parish)
vii-15/21-1971; 19; G. Heinrich. Maryland
Prince Frederick, 7 km S (hardwood forest)
ix-24/xi-14-1987; MT; 19; BRC Hym. Team
Oklahoma: Red Oak (Latimer Co.); ix-1993
FIT; 19; K. Stephan. South Carolina: Harley-
ville, 10 km NE, [and] Harleyville, nr.; 29,
IS; (see Types). Pendleton (225 m); vi-24/
vii-1-1987, viii-6/ 14-1987, ix-5/9-1987; MT;
59; BRC Hym. Team. Pendleton, Tangle-
wood Spring, 34 38.7' N, 82 47.1 'W
(225 m); ix-15/30-1987; MT; 19; J. Morse.

Comments. — There are an additional three
or four closely related species occurring in
Central and South America. A. primum
females is separated by the following com-
bination of traits: midlobe of mesoscutum
with suboval to subrhomboidal reticulations
laterally with cells weakly corrugated with-
in; mandibular teeth regular in shape;
antennal club with subglobose BPS and
spiniform PLS. In the undescribed species,
the mesoscutum either is linearly reticulate,
or with cells which are smooth within. In at
least one species one of the posterior
mandibular teeth is apically bifurcate. In at
least two of the undescribed species the BPS
are narrow and subfusiform in shape.
Finally, in a species from Costa Rica the
female lacks spiniform PLS. Males are
available for more than one of the unde-
scribed species; all have the same antennal
dimorphism indicated for A. primum.

22. Adryas Pinto and Owen

(Figs 67, 68, 140, 141)

Adryas Pinto and Owen 2004. Type species:
Adryas magister Pinto and Owen, by original
designation.

Volume 15, Number 1, 2006

Diagnosis. — Female. Antenna with 1 anel-
lus, and 3-5 club segments but club with
segments variously fused, in total or in
part, and never with 5 completely separat-
ed segments, last segment of club with at
least a few RS ventroapically; CI anelli-
form, very closely appressed to C2. Fore
wing variable but with following charac-
teristics: venation elongate, attaining 0.6-
0.8 FWL, MV and PM confluent, PM
triangular, widest at base, distinctly wider
than MV, deviating minimally if at all from
wing margin, with a distinct gap between
SC and PM; disk fumate behind venation,
with linear setal tracks apical to fumate
area, RSI present or not.

Male. Antennal club completely 5 seg-
mented and without RS in species whose
males are known. GC lacking PAR and VS,
ventral setae present; aedeagal AAP ab-
sent.

Distribution. — Central and South Ameri-
can tropics.

Diversity. — Adryas, with eight species,
was recently described (Pinto and
Owen 2004); none of its species had been
known previously. Included are A. albicer-
ata, A. bochica A. erwini, A. incompta, A.
iris, A. lioptera, A. magister and A. plurifu-
mosa. Although the genus is widespread
in the American tropics with relatively
high levels of sympatry, representatives
are not commonly collected. A key to
species is provided by Pinto and Owen
(2004).

Discussion. — Adryas is distinguished by
antennal formula and wing venation. The
anelliform CI, incompletely separated club
segments in females, and the shape and
position of the premarginal vein are dis-
tinctive. Uscanoidea and Zaga, also with
a five-segmented club and similarly anelli-
form CI are separated by wing venation
characterized by the premarginal which is
narrower or subequal in width to the
marginal vein, never wider. The fumate
fore wing and presence of recurved setae
on the club further separate Adryas from
these genera. Uscanopsis which also may be

confused with Adryas does have a relatively
wide premarginal vein. However, the
shape of this vein is not comparable
(elongate and widest at the middle, not
subtriangular and widest basally as in
Adryas). The elongate apical spur of the
hind tibia in Uscanopsis (c. 0.9 the length of
entire hind tarsus) is a further distinguish-
ing trait.

Records. — Belize. Bolivia. Colombia.
Costa Rica. Ecuador. Peru. Venezuela.
See Pinto and Owen (2004) for detailed
collection records of all eight species.

Hosts. — Unknown.

23. Aphelinoidea Girault

(Figs 46, 121-123, 163, 175, 227, 228)

The synonymy below follows Doutt and Vig-
giani 1968.

Aphelinoidea Girault 1911. Type species: Apheli-
noidea semifuscipennis Girault, by original
designation.

Lathromeroides Girault 1913. Type species: La-
th.romeroid.es longicorpus Girault, by original
designation. Currently as subgenus.

Diaclava Blood and Kryger 1928. Type
species: Diaclava waterhousei Blood and Kry-
ger, by original designation. Currently as
subgenus.

Krygeriola Nowicki 1934. Type species: Kryger-
iola dolichoptera Nowicki, by original designa-
tion. Originally as subgenus of Aphelinoidea;
currently synonym of Lathromeroides.

Thalesanna Girault 1938. Type species: Apheli-
noidea nigrioculae Girault, by original desig-
nation. Currently as subgenus.

Lengerkeniola Nowicki 1946. Type species: Len-
gerkeniola bischoffi Nowicki, by monotypy.
Originally as subgenus of Lathromeroides;
currently synonym of Lathromeroides.

Encyrtogramma De Santis 1957. Type species:
Encyrtogramma habros De Santis, by original
designation. Currently as subgenus.

Tanygramma De Santis 1957. Type species:
Tanygramma xiphias De Santis, by original
designation. Currently synonym of Lathro-
meroides.

Diagnosis. — Female. Antenna with 2 anelli
and an elongate, narrow club of 2 or 3
segments, basal club segment commonly

Journal of Hymenoptera Research

incompletely divided, club with numerous
APB sensilla, both sexes lacking a terminal
process at apex of club. Fore wing 2.1-2.5X
as long as wide, fringe setae length variable
(0.1-0.3 X FWW); venation short, attaining
< 0.4 distance to apex of wing, SV reduced,
either short truncate and sessile to MV or,
less commonly, represented by a slender
appendix projecting from the marginal
vein; disk densely setose, few if any
arranged in linear tracks, RSI absent. Hind
wing with at least 3 complete linear setal
tracks.

Male. GC with aedeagal AAP present or
not; PAR absent, VS present but vestigial,
lobiform; ventral setae present.

Distribution. — Worldwide, apparently
most diverse in temperate areas. Of the
three currently recognized New World
subgenera, the nominate and Lathromeroi-
dea occur throughout the hemisphere;
Encyrtogramma is restricted to South Amer-
ica.

Diversity. — Aphelinoidea includes 36 spe-
cies; nine have been described from the
New World . These are partitioned among
the three subgenera as follows: Lathromer-
oides [neomexicann (Girault) (North Amer-
ica) and xiphias (De Santis) (South Amer-
ica], Encyrtogramma [habros (De Santis) and
semiflava De Santis, both South America],
and the nominate subgenus [oblita De
Santis (South America), plutella Girault,
roja Triapitsyn, Walker and Bayoun, semi-
fuscipennis and zarehi Triapitsyn, Walker
and Bayoun (the latter four described from
North America)]. In addition two Palearc-
tic species of the nominate subgenus
[anatolica Nowicki, turanica Trjapitzin] have
been introduced into North America for
biological control (Trjapitzin 1995). Identi-
fication of species of the nominate sub-
genus remains problematic with several
species undescribed (Trjapitzin 1995). Par-
tial keys to the U.S. species are provided by
Trjapitzin (1995) and Walker et al. (2005).

Discussion. — Aphelinoidea is readily iden-
tified by the relatively narrow and highly
setose fore wing which lacks linear setal

tracks, the reduced stigmal vein, the
absence of a funicle and the two-three
segmented antennal club which lacks
a terminal process. It is most likely con-
fused with Tumidiclava. Fore wing setation
and venation as well as antennal structure
are similar in both (see Tumidiclava for
separation). Aphelinoidea may be more
closely related to the Brachygrammatella
group, although separation is straightfor-
ward owing to the presence of a funicle in
the latter. Similarities include antennae
with numerous APB sensilla on the club,
densely setose fore wings, poorly devel-
oped stigmal vein, and similar male geni-
tala. Interestingly, as in Aphelinoidea, the
first postanellar segment in Chaetogramma
is commonly incompletely divided.

The validity of the five subgenera re-
quires study. For example, limits based on
length of metasoma vs mesosoma for
separating the nominate subgenus from
Lathromeroidea appear to be artificial with
intermediate undescribed species. The
nominate subgenus also is inadequately
defined. Trjapitzin (1995) divided the as-
semblage into two species groups, the
anatolica group which includes the North
American A. plutella, and the monotypic
semifuscipennis group. Walker et al. (2005)
recently changed the name of the anatolica
group to the plutella group which is
characterized by the presence of a hyaline
path on the fore wing immediately apical
to the venation. This modification is absent
in the semifuscipennis group. Male genitalic
variation also separates the two. Thus in
the plutella group aedeagal apodemes are
present, whereas they are absent in the
semifuscipennis group. Apodemes also are
present in Lathromeroidea and Encyrto-
gramma. The latter subgenus differs from
the plutella group only by the distinct fore
wing setation and color pattern; it also has
a hyaline path on the fore wing. This
renders the nominate subgenus paraphy-
letic at best and it appears that the plutella
group should be transferred to Encyrto-
gramma. However, I hesitate to revise the

Volume 15, Number 1, 2006

definition of these taxa until all subgenera
can be considered.

New World records. — Subgenera (A/E/L)
are indicated after each record. Argentina:
Buenos Aires (E/L), Catamarca (A),
Chaco (E), Formosa (A/E), La Pampa (E),
La Rioja (A/E), Mendoza (E), Misiones (A),
Salta (A/E), Tucuman (E). Bermuda: (A).
Brazil: Rio de Janeiro (E), Sao Paulo (A).
Bolivia: (A/E). Canada: Alberta (A/L),
Manitoba (A), Northwest Territories (A),
Ontario (A), Yukon Territory (L). Chile:
Metropolitana (A), Region V (E), Region IX
(A), Region X (A). Costa Rica: (A/L).
Ecuador: (A/L). Mexico (A only): Baja
California Norte, Baja California Sur, Cam-
peche, Chiapas, Colima, Guerrero, Nuevo
Leon, San Luis Potosi, Sinaloa, Tamaulipas,
Veracruz. United States: Alabama (A),
Arizona (A), California (A/L), Colorado
(A), Florida (A), Georgia (A), Illinois (A),
Indiana (A), Kansas (A), Louisiana (A),
Maryland (A), Michigan (A), Missouri (A/
L), Montana (A), Nebraska (A), Nevada (A),
New Mexico (A), North Carolina (A),
Oklahoma (A/L), Oregon (A/L), South
Carolina (A), Texas (A/L), Utah (A/L),
Virginia (A), Washington (A), Wyoming
(A/L). Uruguay: (A/E/L). West Indies: (A).

Hosts. — Hemiptera: Cicadellidae (Trja-
pitzin 1995, Walker et al. 2005), Fulgoridae
(Timberlake 1924).

24. Bloodiella Nowicki

Bloodiella Nowicki 1935. Type species: Blood-
iella andalusica Nowicki, by original designa-
tion.

Diagnosis. — Female. Antenna with 1 anel-
lus, 1 funicular and 3 club segments. Eyes
reddish. Maxillary palp narrow, elongate,
c. 0.8X mandibular length. Midlobe of
mesoscutum and scutellum each with 2
pair of elongate setae. Fore wing very
broad, c. 1.6X as long as wide; disk
moderately densely setose with setae ar-
ranged in linear tracks, RSI present. Meta-
somal tergum I (petiolar) without a trans-
verse row of denticles.

Male. GC without aedeagal AAP, VS or
PAR (Viggiani 1971, Fig. 5).

Distribution. — Palearctic, Africa?, South
America?.

Discussion. — There are three species now
assigned to Bloodiella, however the diagnosis
above is based only on the type species, B.
andalusica. This is because the other two are
questionably placed. In the New World only
B. carbonelli De Santis is included. B.
onnenidis (Dozier), provisionally moved to
Bloodiella (Doutt and Viggiani 1968), is now
transferred to Burksiella (see below). The
latter authors also questioned generic place-
ment of the African species, Bloodiella
gynandroplithalniae (Risbec). The assignment
of carbonelli to Bloodiella may also be in error.
Its unique holotype (in MLPA) was exam-
ined but because of the poor mount I am
unable to verify number of funicle segments.
It is possible that a small anelliform Fl can
not be seen in the type, which, if present,
also would require transfer to Burksiella.

Bloodiella is one of only three genera
other than representatives of the Oligositini
with one funicle and three club segments.
The other two are Soikiella, and the African
Oligositoides. Only Soikiella occurs in the
New World. Unlike Bloodiella, Soikiella
lacks an RSI setal track, has a transverse
row of denticles on metasomal tergum I,
and has short aedeagal apodemes as well as
parameres and volsellae in males. From the
Oligositini, these three genera are separated
by several features including the much
broader fore wings, reddish eyes, and the
two pair of setae on both the midlobe of the
mesoscutum and the scutellum.

New World records. — Uruguay (holotype
of B. carbonelli).

Hosts. — Unknown.

25. Brachista Walker

(Figs 47, 124, 265)

Brachista Walker 1851. Type species: Brachysticha
fidiae Ashmead, by subsequent monotypy.
Pinto 1994 (revision).

Brachysticha Forster 1856. Unjustified emenda-
tion.

Journal of Hymenoptera Research

Giraultiola Nowicki 1936. Type species: Brachys-
ticha fidiae Ashmead, by monotypy. Pinto
1993 (synonymy).

Diagnosis. — Body elongate, relatively
narrow. Female. Antenna with 2 anelli and
a 4 segmented club, CI as long or longer
than C2 and bearing 1 PLS. Fore wing 2.1-
2.5 X as long as wide, fringe setae variable
in length, 0.1-0.4 X FWW; venation with SV
well developed; disk moderately densely
setose, setae relatively short and not
obviously arranged in linear tracks, a dis-
tinct RSI absent. Metasoma with 2 elongate
ventrolateral strut-like apodemes extend-
ing anteriorly from near base of ovipositor
(Fig. 265).

Male. Without strut-like metasomal apo-
demes. GC tubiform with a relatively
elongate anterodorsal aperture; PAR, VS,
ventral setae, and aedeagal AAP absent.

Distribution. — Known from the New
World only, from Canada to Costa Rica,
and SW Brazil.

Diversity. — Three species described: B.
fidiae, B. efferiae Pinto, B. fisheri Pinto.

Discussion. — Brachista is most similar to
Uscana. In both the antennal club is four
segmented, and a funicle is absent. Also, in
both, CI is relatively elongate and bears at
least one PLS. The two are separated by
fore wing setation (linear tracks including
RSI present in Uscana; distinct tracks with
RSI absent in Brachista), and the absence of
strut-like apodemes at the apex of the
metasoma in Uscana females. The poorly
known Lathromeromyia from Asia may also
be close to Brachista; its females lack the
characteristic metasomal apodemes as
well. Brachista was studied by Pinto
(1994). The rather tortuous history of its
generic nomenclature also was reviewed
(Pinto 1993).

Records. — Brazil: Rondonia. Canada:
Quebec. Costa Rica. Guatemala. Mexico:
Baja California Sur, Chiapas, Guerrero,
Hidalgo, Jalisco, Michoacan, Morelos,
Nayarit, Oaxaca, Sinaloa, Tamaulipas.
United States: Arizona, California, Illinois,

Maryland, North Carolina, Ohio, Pennsyl-
vania, Texas, Virginia. West Indies.

Hosts. — Coleoptera: Chrysomelidae
(Ashmead 1894). Hemiptera: Cicadidae
(Marlatt 1898). These records are for 8.
fidiae. The other two species of Brachista are
phoretic on Asilidae which may also serve
as hosts (Pinto 1994).

26. Brachygrammatella Girault

(Fig. 125)

Brachygrammatella Girault 1915. Type species:
Brachygrammatella nebulosa Girault, by origi-
nal designation. Doutt 1968 (generic review).

Pseudbrachygramma Girault 1915. Type species:
Pseudbrachygramma perplexa Girault, by orig-
inal designation. Doutt and Viggiani 1968
(synonymy; as subgenus, currently valid).

Diagnosis. — Female. Antenna with 2 anelli,
2 funicular and 1 or 2 club segments (often
incompletely divided), Fl and F2 transverse,
closely appressed but similar in length and
width; PLS present on funicle, numerous
short, fine APB sensilla on club. Fore wing
wide, <2X as long as wide, fringe setae very
short, < 0.1 X FWW; venation with base of
MV and apex of PM poorly sclerotized, light
in color, SV short, lacking a basal constric-
tion, MV much broader than PM with
numerous heavy setae dorsally; disk dense-
ly setose with considerable setation extend-
ing basally behind venation, distinct linear
setal tracks absent or few in number, RSI
absent, costal cell well developed due to
narrow PM. Ovipositor not extending be-
yond apex of metasoma.

Male. GC with vestigial, lobiform VS,
ventral setae and aedeagal AAP present.

Distribution. — Australia, Asia and Africa.
A single record from Mexico.

Diversity. — There are 11 species of Bra-
chygrammatella; none is known from the
New World. A single collection of two
females from Mexico (UCRC), the only
New World record, has not been identified.

Discussion. — Two subgenera currently
are recognized (Doutt 1968). The characters
used for subgeneric separation fail for

Volume 15, Number 1, 2006 91

certain apparently undescribed species in ward base of wing, apex of RSI converging

the UCRC. The Mexican specimens how- on Cu tracks.

ever, are assignable to the subgenus Pseud- Male. Antenna with club segments usu-

brachygramma. ally less asymmetrical; benefica group males

Brachygrammatella is closely related to with several APB on CI. GC reduced,

Chaetogramma and Xiphogramma. Differ- gradually narrowed to apex, basal margin

entiating characters and a discussion of rounded; with or, more commonly, with-

generic limits are treated under Chaeto- out ventral setae, PAR and aedeagal AAP;

gramma. GL considerably less than HTL.

New World records.— Mexico: Sinaloa. Distribution.— Oriental Region and New

Hosts.— Hemiptera: Cicadellidae (Doutt World. In the New World Burksiella is

1968), Membracidae (Viggiani and Hayat broadly distributed from Canada S to

1974), Miridae (Viggiani 1968). Argentina and Chile. Unlike the similar

genus Zagella, it is most diverse in tropical

27. Burksiella De Santis, renewed status areas where virtually all species are un-

(Figs 69-71, 142, 161, 242, 243, 266-274) described.

_ , . „ _ _ incr7 „ D / n Diversity. — There are nine species as-

Burksiella De Santis 1957. Type species: Burksiella . , , y n , . „ . , - r. _

, , L „ c v • . i j • signed to Burksiella (see below). Seven

subannulata De Santis, by original designa- & \

occur in the New World and two are
hon.

Parahispidaphila Yousuf and Shafee 1988: 136. Asian. Numerous undescribed species ex-
Type species: Varahispidophila singularis You- ist; only one of these, Burksiella dianae from
suf and Shafee, by original designation. New North America, is described here.
synonymy. Discussion. — Burksiella has been treated

as a junior synonym of Zagella (Doutt and

Diagnosis.— Female. Antenna with 2 an- viggiani 1964) and it is that genus which is

elli, 2 funicular and 3 club segments (rarely most \^eiy to be confused with it. Fore

reduced to 2 with only a partial division wjng structure provides the easiest means

between CI and C2); F2 as wide as or, more or separation. In Burksiella (Figs 142, 161)

commonly, wider than CI, usually with the stigmal vein usually is constricted

more than 1 PLS; Fl short, transverse, basally and always is projected apically

closely appressed to F2; club segments beyond the end of the marginal vein. In

distinctly asymmetrical (i.e., length of CI Zagella (Figs 150, 162) on the other hand,

and C2 in particular varying with surface), the stigmal vein is not constricted and is

CI without or, uncommonly (benefica perpendicular to the marginal vein rather

group), with a single APB. Fore wing 1.7- than projecting apically. The length and

1.9X as long as wide, fringe setae short (< orientation of the RSI provide another

0.1 X FWW); venation usually with a con- difference. In Burksiella it is longer and

striction between stigma and MV, SV curved posteriorly toward the base of the

extending apicad of apex of MV (a line wing and converges onto the Cu tracks at

drawn through middle of SV to wing its apex. In Zagella the RSI has fewer setae

margin describes an oblique angle with and is relatively straight and roughly

MV), MV 1.3-1.5 X as long as PM and 0.9- perpendicular to the Cu tracks. The longer,

1.7X as wide, SV+MV/PM > 1.5 (usually narrower marginal vein coupled with the

> 1.75), PM and base of MV not distinctly greater extension of the stigmal vein

less sclerotized than apex of MV, radial apically also allows use of the ratio MV +

process present; disk moderately densely SV/PM for generic separation. Antennal

setose, with distinct linear setal tracks, RSI characters may also be useful. In most

track elongate with more than 5 setae and Burksiella CI lacks APB sensilla and F2 is as

broadly curved from apex of stigma to- wide or wider than the club; in Zagella APB

Journal of Hymenoptera Research

sensilla are always present on CI and F2 is
slightly narrower than the club. Males
usually can be distinguished by genitalia.
In Burksiella, the genital capsule gradually
narrows to its apex and its basal margin is
typically arcuate (Figs 242, 243); in Zagella
the capsule more abruptly narrows near
the middle and the basal margin is trans-
verse or emarginate (Figs 254-256).

Although most likely confused with
Zagella because of similar antennal formu-
la, Burksiella appears more closely related
to Uscanoidea. Uscanoidea is defined as
having a five-segmented club but in certain
species the distinction between funicle and
club is not straightforward. Wing and male
genitalia are similar in both groups and I
have not discovered differences other than
antennal formula.

As in certain other genera male genitalic
variation in Burksiella is considerable. Most
species are as in B. spirit a. The genital
capsule is simple, lacking parameres, ven-
tral setae and aedeagal apodemes. In
others such as B. florida and certain
undescribed Latin American species, how-
ever, these structures are present.

Numerous New World species of Burk-
siella await description. The only species
named are Burksiella subannulata, B. dianae
n. sp. (described below), B. florida (Vig-
giani) new combination (from Zagella), B.
spirita (Girault) new combination (from
Zagella), B. plateusis (De Santis) new com-
bination (from Monorthochaeta), B. orme-
nides (Dozier) new combination (from
Bloodiella) and B. benefiea (Dozier) new
combination (from Ufens). The type mate-
rial of all species was examined. It is
possible that Bloodiella carbonelli De Santis,
from Uruguay, belongs here as well but I
am treating it under the original combina-
tion for now (see Bloodiella). Burksiella
ormenides, originally described in Ufens by
Dozier (1932), was provisionally trans-
ferred to Bloodiella by Doutt and Viggiani
(1968). Examination of the type material of
this species (NMNH) clearly shows two
funicular segments, not one as is charac-

teristic of Bloodiella. All other traits support
transfer to Burksiella as well.

Species of Burksiella occur in Asia. In-
cluded is B. chrysomeliphila (Lin) new
combination (from Zagella) and B. singu-
lars (Yousuf and Shaffee) new combina-
tion (from Parahispidoplula). It should be
noted that the illustration of the latter
species in Yousuf and Shafee (1988,
Fig. 24F) is misleading. It shows the two
funicular segments as subequal in length.
The holotype was examined (BMNH) and
it is clear that Fl is much shorter than F2 as
is typical of Burksiella and related genera.

Burksiella as defined here includes what
can be referred to as the benefiea group.
This assemblage differs considerably from
other species in that the sexes are highly
dimorphic. Females resemble congeners,
but as noted by Dozier (1932) in his
description of benefiea, males appear as
a different species. They are larger, much
lighter in color, and megacephalic. This
group also differs in that CI bears a single
strong APB sensillum in females and males
have several APB on this segment. APB
sensilla are almost never present on CI of
other Burksiella. Although the West Indian
B. benefiea is the only species of the group
described, two or three additional North
American species are known. They are
parasitoids of tettigoniid eggs.

New World records. — Argentina: Buenos
Aires, Catamarca, Chaco, Cordoba, For-
mosa, Jujuy, La Rioja, Misiones, Salta, San
Luis, Tucuman. Belize. Bolivia. Brazil:
Guanabara, Mato Grosso do Sul, Minas
Gerais, Rondonia, Sao Paulo. Canada:
Alberta, Ontario, Quebec. Chile: Regions
IX, X. Colombia. Costa Rica. Ecuador.
Guatemala. Mexico: Aguascalientes, Baja
California Sur, Campeche, Chiapas, Co-
lima, Guerrero, Jalisco, Michoacan, Mor-
elos, Nayarit, Nuevo Leon, Oaxaca, Quin-
tana Roo, San Luis Potosi, Sinaloa, Tamau-
lipas, Veracruz, Yucatan. Nicaragua. Peru.
United States: Arizona, California, Florida,
Georgia, Illinois, Kansas, Louisiana, Mary-
land, Mississippi, Missouri, Nebraska,

Volume 15, Number 1, 2006

New Hampshire, Nevada, North Carolina,
Oklahoma, Oregon, South Carolina, South
Dakota, Texas, Utah, Washington, Wyom-
ing. Venezuela. West Indies.

Hosts. — Coleoptera: Chrysomelidae (Lin
1994). Hemiptera: Ckadellidae (Triapitsyn
et al. 1998 and Triapitsyn 2003 [as Zagella in
both], unpubl. records). Orthoptera: Tetti-
goniidae (Dozier 1932, unpubl. records).

Burksiella dianae Pinto, new species

(Figs 266-274)

Description. — Body relatively compact,
robust, length (card-mounted specimens)
0.4-0.5 mm; head broad, c. 1.2X greatest
mesosomal width. Color dark brown ex-
cept top and front of head, antennae and
usually very base of metasoma yellow
brown, metanotum usually white at mid-
dle; legs lighter at junction of femur and
tibia; tarsi much lighter. Female. Mandible
with 3 teeth, the two posterior teeth large,
distinct, the anterior tooth short, obsoles-
cent. Maxillary palp elongate, widest at
base, c. 5x as long as wide, without a peg-
like sensillum apically, terminal seta slight-
ly longer than palp, subapical seta c. 2/3
palpal length. Antenna with 2 discoid
anelli, funicle distinctly transverse and
somewhat swollen, club subcorneal and 3
segmented; length/width of scape, pedicel,
funicle and club: 43/15, 24/21, 27/34, 72/
27; Fl closely appressed to F2, visible in
medial view only; club widest at base, CI
and C2 asymmetrical, length/width of
segments: 27/25, 40/30, 30/13; sensilla:
BPS small, mushroom shaped, present on
all postanellar segments, PLS on F2-C3 (4
on F2), APB on Fl, F2 and near apex of C3,
FS on all club segments, longest FS
subequal to greatest club width, UPP at
apex of club setiform.

Midlobe of mesoscutum elongate, 1.2X
as long as wide; midlobe and scutellum
each with 2 pair of moderately long setae,
posterior pair on midlobe slightly more
medially placed than anterior pair; sculp-
turing on midlobe and scutellum closely

lineate with several transverse interlineal
connections on midlobe, relatively few on
scutellum. Propodeum narrow, discal area
only slightly produced posteriorly, sub-
equal in length to metanotum. Fore wing
broad, oblate, 1.6X as long as wide,
venation attaining 0.5 FWL, fringe setae
very short, c. 0.05 X FWW; venation robust,
MV c. 1.4X length of PM, gradually in-
creasing in width to apex, c. 2.5 X as long as
wide, with a relatively large gap separating
MV and PM; SV c. 3A length of MV, stigma
suboval, SV+MV/PM = 1.6-1.7; MV with 3
moderately long anterior and 4 shorter
posterior setae dorsally, with 4 ventral
setae; PM with 2 elongate setae and 2
campaniform sensilla at apex, PM consid-
erably behind anterior wing margin
forming a relatively large costal cell
with several setae on its anterior margin
and 1 or 2 posterior setae; disk moderately
densely setose, almost all setae associated
with linear tracks (c. 19 in number),
RSI present, broadly curved, basal track
with 2 setae. Hind wing with 3 linear
setal tracks. Metasomal terga lineate. Ovi-
positor relatively short, robust, OL/HTL =
0.8-1.0, gonoplac short, sharply and asym-
metrically pointed at apex; hypopygium
short, subtending only basal third of
ovipositor.

Male. Antenna less robust than in female,
club smaller, with CI and C2 partially
fused and with fewer sensilla. Metasoma
distinctly swollen apically, distinctly bi-
lobed. GC short, GL slightly less than half
HTL, middle of basal margin with a short
cuticular flange projecting anteriorly.

Types.— Holotype 9 and allotype ;. UNITED
STATES. California: Riverside Co., Menifee
Valley (hills on W side), 33 19' N, 117 13'VV
(1800' el.); vii-31-1995 (holotype), vii-12-1995
(allotype); Malaise trap; J. D. Pinto, collr.; in
NMNH.

Paratypes: 29 with same data as holotype; 1 [
same data except vi-14/28-1995; 2 J same except
x-16/23-1982, xi-1/5-1981, 'pan trap under
Eriogonum gracile'. Paratypes in CNC (1 J, 19 )
andUCRC(2c?/ 19)

Journal of Hymenoptera Research

Etymology. — After my wife, Diana Gail
Pinto.

Distribution. — United States and Mexico;
from Washington state E to Illinois, S to
southern Mexico.

Comments. — Burksiella dianae is unique. It
is easily separated from B. spirita, the only
other named North American Burksiella, as
well as from all other species, described
and undescribed. Both sexes are distin-
guished by the maxillary palp. In all other
species the palp is shorter, but more
significantly, bears on its apex in addition
to a terminal seta, a distinct digitiform or
peg-like sensilla (as in Fig. 172). In B.
dianae, the elongate and tapered palp has
a terminal seta only (Fig. 269). Males of B.
dianae are distinguished from other species
by the apically swollen and bilobed meta-
soma (Fig. 273). The dorsolateral metaso-
mal surface in this area is membranous and
appears to be densely porose. The short but
distinct flange at the base of the genital
capsule also appears to be unique (Fig. 274).
The transverse funicle in females also helps
separate this species from most other
Burksiella where this segment is more
elongate.

Material examined. — 469 (44 on slides), 16 ^ (15
on slides). MEXICO. Colima: Manzanillo; viii-8-
1984; 19; G. Gordh. Guerrero: Acapulco, 30 km
N; viii-6-1984; SP; LJ; G. Gordh. Michoacan:
Lazaro Cardenas, 100 km N; viii-7-1984; lo; G.
Gordh. Oaxaca: Yagul Ruins; vii-13-1984; G.
Gordh. Sinaloa: La Concha, 11 mi. N; x-25-
1982; SP; U; A. Gonzalez. Veracruz: Nautla,
33 mi. S; x-31-1982; 29; A. Gonzalez/J. Huber.
UNITED STATES. Arizona: Bisbee, near; viii-27-
1982; 19; J. LaSalle. Dragoon Mts. (Jordan Cyn.,
Cochise Co.); 31 59' 33" N, 110 01' 07"W; viii-
11-2001; SP; 19; A. Owen. California: Altadena;
vi-9-1991, vi-28/viii-25-1990; 39; R. Crandall.
Anza Borrego State Park (Coyote Cyn., San
Diego Co.); iv-15-1981; lo; J. Huber. Baker,
5.6 mi. SW (at Zzyzx Rd. exit off of 1-15); iii-
30-1989; 'sweeping Larrea, Bebbia, etc.'; 1$; J.
Pinto. Big Pine, 1.1 mi. E (3950' el.); 37 10.39'N,
118° 16.28'W; v-26-2002; 'swp. primarily Salix';
19; J. Pinto. Big Pine, 2 mi. E; vii-12-1993; 'swp.
Salix, etc. along Owens River'; 29; J. Pinto.

Menifee Valley (hills on W. end); 33 39' N, 117
13'W; vi-7/13-1995, x-6/11-1981, x-16/23-1982,
xi-1/8-1982; MT, 'pan trap under Eriogonum';
2 3, 39; J. Pinto. Same locality; 39, 4^ (see Types).
Pozo, 6 mi. SE (1500'); iv-l/v-4-1990; 19; MT; W.
Wahl. Santa Rosa Plateau Ecological Reserve
(590 m) (Riverside Co.); 33° 52.524'N, 117
14.644'W; iv-7/iv-28-2002, iv-29/v-29-2002 ix-
20/X-12-2001, xi-1/22-2001; MT #2; 49, 2&, J.
Pinto. Spanish Flat, 3 mi. SSW on Capell Creek
(Napa Co.); vi-1 1-1984; SP; 19; J. Pinto. Wild-
horse Mdw. (San Bernardino National Forest,
San Bernardino Co.) (2357 m); 34 10.47'N, 116
44.48'W; viii-7-2002; 'yellow pans riparian, dry
scrub'; 19, 1 S; A. Owen/J. Pinto. Nevada: Valley
of Fire State Park (1 mi. west of W entrance); iii-
20-1989; 'sweeping flowering plants'; 19; J.
Pinto. Florida: Everglades N. P. (Lone Pine Key
pinelands); vi-6/viii-26-1986; 19; S./J. Peck.
Hollywood; xii-22-1982; sweep; 19; W. Gregory.
Illinois: Garden of the Gods (Harden Co.); vi-19-
1991; 'swp. wooded area'; IS; ]■ Pinto. Oregon:
Christmas Valley, 13.5 mi. SW; viii-5-1995; 'swp.
1 Chrysotluvn nus'; 19; J. Pinto. La Pine, 24 mi SE
(jet. Hwy 35 & Rd. 3125); viii-5-1995; 'swp.
Chrysothamnus etc, in pine forest'; 39; J. Pinto.
South Carolina: Lakeshore, near; vii-25/31-1987;
FIT; 19; J. Johnson. Texas: Ben Bolt, 8 mi. W
(Copita Research Station); v-20-1987, ix-28/30;
49, let; J. Woolley. Big Bend National Park;
(Rosillos Mts.); vii-17-1991; 19; R. Vogtsberger.
Big Bend Ranch SNA (Presidio Co.), McGuirk's
tank (4360'); 29 28' 34"N, 103° 49' 12"W; vi-19-
1990; 19; J. Woolley. Big Bend Ranch SNA
(4400'); 29 ; 30' 45"N, 103 51' 56"W; vi-21-1990;
39, IS; J. Woolley/G. Zolnerowich. Same
except: 29 30' 20"N, 103° 52' 20"W; vi-22-1990;
I9. Washington: Goldendale; vii-21-1988;
'sweep poplar, willow, etc.'; 19; J. Pinto. Wyom-
ing: Rawlings, 17 mi. E (1.5 mi N of 1-80 at
North Platte River); vii-7/ix-2-1991; MT; 29; S.
Shaw.

28. Centrobiopsis Girault, renewed status

(Figs 79, 80, 148, 251)

Centrobiopsis Girault 1918. Type species: Centro-
bia odonatae Ashmead, by original designa-
tion. Doutt and Viggiani 1968 (as synonym of
Lathromeroidea).

Diagnosis. — Female. Body elongate, slen-
der, ovipositor extending considerably be-
yond apex of metasoma. Antenna with 2

Volume 15, Number 1, 2006

anelli and 5 club segments, CI anelliform,
closely appressed to C2, C2 loosely associ-
ated with C3 (may appear as a funicular
segment). Fore wing c. 2x as long as wide,
fringe setae 0.1 X FWW; venation attaining
0.5 FWL, SV constricted at base and
extending apicad of MV, MV 1.5X length
of PM and subequal in width, SV+MV/PM
= 1.8; disk moderately densely setose, with
distinct linear setal tracks, RSI elongate,
broadly curved from apex of stigma toward
base of wing, its apex converging on
Cu tracks. Propodeal disk length subequal
to that of metanotum. Fore tibia not spined
on dorsal surface. Ovipositor elongate,
extending entire length of metasoma, c. lA
its length extending beyond metasomal
apex.

Male. GC elongate, slender, its length c.
1.6 X HTL, with an elongate, longitudinal
furrow on ventral surface; PAR, VS, ventral
setae and aedeagal AAP absent.

Distribution. — North America.

Diversity. — The only named species is
Centrobiopsis odonatae, renewed combina-
tion (from Lathromeroidea). One additional
undescribed species has been collected
from southern Mexico.

Discussion. — Centrobiopsis is not highly
distinctive yet it is not easily placed in any
other genus. Its body and genitalia are
elongate, with both the male genital cap-
sule and ovipositor considerably longer
than the hind tibia. In these respects it
resembles Chaetostricha. Unlike Chaetostri-
cha, however, a distinct funicle is absent,
the fore tibia is not spinose, and although
the male genital capsule is narrow and
exceptionally elongate in both, only in
Centrobiopsis is the ventral surface fur-
rowed (cf. Figs 245, 251).

Doutt and Viggiani (1968) synonymized
Centrobiopsis with Lathromeroidea. However,
unlike Lathromeroidea, in Centrobiopsis the
fore wing fringe is very short, it lacks the
conspicuous basal convergence of setal
tracks characteristic of Lathromeroidea in
the original sense, and unlike most species
of that genus, its propodeal disk is sub-

equal in length to that of the metanotum.
The unique male genitalia further distances
it from Lathromeroidea and all other genera
of the Chaetostricha group. Affinity seems
closer to Uscanoidea, which shares most of
the fore wing characteristics as well as the
shorter propodeal disk. Although similar
to both genera as well as to Burksiella, C.
odonatae is not easily placed in any one of
them without considerably modifying ge-
neric definitions.

Ashmead's type of C. odonatae (exam-
ined, NMNH) is incomplete. Only a fore
and hind wing, and one antenna remain.

Records. — Canada: Ontario, Quebec.
Mexico: Quintana Roo. United States:
Illinois.

Hosts. — Odonata: Lestidae (Ashmead
1900, several unpubl. records).

29. Chaetogramma Doutt

(Figs 48, 49, 126, 193, 229, 230)

Chaetogramma Doutt 1974. Type species: Chaeto-
gramma occidentalis Doutt, by original desig-
nation.

Chaetogrammina Hayat 1981 (as subgenus, cur-
rently valid). Type species: Chaetogramma
(Chaetogrammina) maculata Hayat, by original
designation.

Brachistagrapha De Santis 1997. Type species:
Brachistagrapha caudata De Santis, by original
designation. Pinto and Viggiani 2004 (synon-
ymy).

Diagnosis. — Female. Antenna with 2 an-
elli, funicle and 1 club segment, funicle
consisting of 2 subequal closely appressed
segments or a single partially divided
segment; PLS present on funicle, with
numerous short, fine APB sensilla on club.
Fore wing venation with base of MV and
apex of PM poorly sclerotized, light in
color, SV short, lacking a basal constriction,
MV without numerous heavy setae dorsal-
ly; disk densely setose but dense setation
not extending basally to area behind
venation, with linear setal tracks poorly
indicated, RSI absent. Ovipositor shorter
than metasoma and not extending beyond
its apex.

Journal of Hymenoptera Research

Male. GC with vestigial, lobiform VS,
ventral setae present; aedeagal AAP pres-
ent or not.

Distribution. — Asia, Africa, North and
temperate South America.

Diversity. — Six species are assigned cur-
rently to Chaetogramma. Only two occur in
the New World, Chaetogramma occidentalis
and C. caudata (De Santis). Both belong to
the nominate subgenus. The latter species
was designated as the type of Brachistagra-
pha by De Santis (1997) who erroneously
assigned it to the Oligositini (Pinto and
Viggiani 2004).

Discussion. — Chaetogramma is closely re-
lated to Brachygrammatella and Xipho-
gramma. Generic limits in this complex ( =
Brachygrammatella group) require clarifica-
tion. In general, Brachygrammatella is de-
fined by the broad, dorsally spinose mar-
ginal vein (Fig. 125), the larger costal cell
(due to reduction of PM width), the trans-
verse funicular segments, and the basal
extension of the dense discal setation of the
fore wing into the region behind the
venation. Brachygrammatella is the only
genus in the group that may have a two-
segmented club. Xiphogramma is character-
ized by the absence of Brachygrammatella
features and its elongate ovipositor which
extends considerably beyond the apex of
the metasoma (Fig. 195). Chaetogramma is
distinguished by the absence of the pre-
sumably derived features characterizing
Brachygrammatella and Xiphogramma. In
Chaetogramma the funicle is two segmented
with Fl and F2 partially (C. Chaetogramma)
(Figs 48, 49) or completely (C. Chaetogram-
mina) divided. Male genitalia are similar in
all three genera. Although aedeagal apo-
demes are present or absent in this group,
variation does not coincide with generic
limits. Thus, apodemes are absent in
Xiphogramma fuscum (Fig. 239) and C.
maculata, but present in X. anneckei and C.
occidentalis (Fig. 229); they also are present
in Brachygrammatella (Viggiani 1971). Males
are unknown for X. indicum Hayat and X.
holorhoptra Nowicki.

Hayat (1981) recognized the Asian sub-
genus Chaetogrammina primarily by the
completely divided funicle, discernable
setal tracks basally in the fore wing, and
a better developed costal cell. As indicated
by Pinto (1990b) these traits also character-
ize Xiphogramma. The only difference be-
tween Chaetogramma and Xiphogramma is
the longer ovipositor in the latter.

Intermediate states and lack of consis-
tency in features used by Doutt (1968, 1973)
and Doutt and Viggiani (1968) to distin-
guish Brachygrammatella, Xiphogramma and
Chaetogramma render these genera difficult
to define on a world-wide basis (Pinto
1990b). Studies of material from Asia,
Africa and Australia, where the bulk of
the species occurs, is needed. Problematic
generic limits currently do not affect the
New World fauna from a practical stand
point where only two closely related
species of Chaetogramma, and one species
of Xiphogramma and of Brachygrammatella
are known.

New World records. — Argentina: La Rioja,
Salta, Misiones. Canada: Alberta, Manitoba,
New Brunswick, Ontario, Prince Edward
Island. Chile: Region X (Isla Chiloe). Costa
Rica. Mexico: Baja California Norte, Co-
lima, Guerrero, Jalisco, Morelos, Nuevo
Leon, Sinaloa, Tamaulipas, Yucatan. United
States: Arizona, California, Colorado,
Louisiana, Montana, Oregon, Texas, Virgi-
nia, Washington. West Indies.

Hosts. — Unknown.

30. Cliaetostricha Walker

(Figs 72, 143, 244, 245)

Cliaetostricha Walker 1851. Type species: Cliae-
tostricha dimidiata Walker, by monotypy.

Centrobia Forster 1856. Type species: Tricho-
gramma walkeri Forster, by monotypy.

Centrobiella Girault 1912. Type species: Centro-
biella mulierum Girault, by original designation.

Ratzeburgalla Girault 1938. Type species: Paru-
fens spinosus Girault, by original designation.

Diagnosis. — Female. Body usually slender
and elongate; head relatively narrow,

Volume 15, Number 1, 2006

narrower than mesosoma; ovipositor usu-
ally extending beyond metasoma. Antenna
with 2 anelli, 2 funicular and 3 club
segments; Fl short, transverse, closely
appressed to considerably longer F2; F2
longer than wide to as wide as long,
narrower than CI (uncommonly subequal
in width), with 1 PLS (rarely absent); club
segments relatively symmetrical, almost
always without APB sensilla on CI. Fore
wing 1.8-2.1 X as long as wide, fringe
length 0.08-0.20 X FWW; venation elon-
gate, attaining at least 0.5 FWL, MV
elongate, > 1.2X length of PM; SV+MV/
PM > 1.5 (1.5-2.3), SV constricted at base,
extending apical of MV, stigma often
suboval; disk moderately densely setose
with distinct linear setal tracks, RSI elon-
gate, usually straight and directed toward
wing base. Fore tibia with a prominent
spine at middle of dorsal surface (absent in
some Australian species), if more than one
spine present then middle spine most
prominent.

Male. GC narrow, tubular, usually ex-
tremely elongate, GL subequal to or greater
than HTL, not furrowed ventrally, without
VS, PAR, ventral setae and aedeagal AAP.

Distribution. — Widespread except un-
known from Central and South America.

Diversity. — The genus includes 21 spe-
cies. Only C. thanotophora Pinto, has been
described from the New World but several
additional undescribed forms occur (Pinto
1990a).

Discussion. — As already noted Chaetostri-
cha is superficially similar to Centrobiopsis.
Body shape, ovipositor length and male
genitalia are similar in both (see Centrobiop-
sis for differences). The relatively slender
body shape of Chaetostricha separates it
from similar genera such as Burksiella and
Uscanoidea. They can also be separated by
the fore wing features typical of their genus
and by the genital capsule length which is
considerably less than that of the hind tibia.
Although the dorsum of the fore tibia may
be spinose in a few Uscanoidea, the absence

of a single prominent spine at the middle
also separates it from Chaetostricha.

The synonymy of Centrobia, Centrobielln,
and Ratzeburgalla with Chaetostricha pro-
posed by Doutt and Viggiani (1968) is
followed here. I examined the types of
Centrobielln and Ratzeburgalla (QM) more
than 15 years ago. The types of both are in
poor condition and, although consistent with
the definition of Chaetostricha, I was unable to
verify certain defining characteristics (e.g.
presence of two funicular segments).

New World records. — Canada: British Co-
lumbia, Ontario, Quebec. Mexico: Guer-
rero, Morelos. United States: Arizona,
California, Florida, Michigan, Missouri,
Nevada, Oregon, Texas, Utah, Washington.
West Indies.

Hosts. — Hemiptera: Membracidae (Sri-
vastava et al. 1970), Miridae (Pinto 1990a,
Sundararaju 1993).

31. Lathrotneris Forster
(Figs 50, 127, 231, 262-264)

Lathrotneris Forster 1856. Type species: Lathro-
meris scutellaris Forster, by monotypy.

Lathromerella Girault 1912. Type species: Lathro-
merella fasciata Girault, by original designa-
tion.

Garouella Risbec 1956. Type species: Garouella
ovicida Risbec, by monotypy.

Diagnosis. — Female. Antenna with 2 anelli
and a 5-segmented club, C5 ending in
a pronounced terminal process. Fore wing
2. 1-2.5 X as long as wide, fringe setae 0.2-
0.3X FWW; venation with an elongate,
straight MV and a distinct SV; disk
moderately densely setose, linear tracks
present but indistinct, RSI absent or rarely
poorly indicated. Hypopygium variable
but often prolonged as a narrow projection
extending beneath ovipositor.

Male. Antenna without a terminal pro-
cess. Metasomal terga VI and VII with
a modified area on either side of midline.
GC with PAR present or not, ventral setae
present, VS apparently absent; aedeagal
AAP present.

Journal of Hymenoptera Research

Distribution. — Known from all conti-
nents. In the New World the genus is
known only from the United States, Costa
Rica, Canada and Argentina. It appears to
be uncommon in the neotropics.

Diversity. — There are 23 species assigned
to Lathromeris; most occur in Australia,
Asia and Europe. Only L. argentina De
Santis, a South American species, has been
described from the New World. A distinc-
tive North American representative, L.
hesperus, is described below. Several addi-
tional species, apparently also new, are
known from North America. Certain Aus-
tralian species may be inappropriately
placed in Lathromeris (see below).

Discussion. — Lathromeris is most likely
confused with Tumidiclava and Nicolavespa.
All three have a pronounced terminal
process on the female antennal club. Male
genitalia also are comparable in these
genera, and Tumidiclava males, as in those
of Lathromeris, have similarly modified
metasomal terga. Tumidiclava is separated
by its extremely short stigmal vein which
lacks a constriction between the stigma and
marginal vein; also females of Tumidiclava
have fewer than five antennal club segments
and males frequently do as well. Nicolavespa
has a distinct funicle and its males lack
modifications of the metasomal terga.

It is likely that certain Australian species
assigned here are misplaced. Girault (1912)
defined Lathromerella, a junior synonym of
Lathromeris, as having a five-segmented
club and considered the terminal process
at its apex in females to vary within the
genus (e.g. Girault 1915). He erroneously
considered Pterygogramma as having
a three-segmented club (Girault 1912).
The club is five-segmented in both genera
but the terminal process is lacking only in
Pterygogramma. Consequently it appears
that Girault placed certain species of
Pterygogramma in Lathromerella if the five-
segmented club was detected. Species
have been misplaced include L.
It, L. occidentalis Girault and L.
ovid ilt.

Neiv World records. — Argentina: Buenos
Aires (De Santis 1957). Canada: Alberta,
British Columbia, Ontario, Prince Edward
Island, Quebec. Costa Rica. United States:
Arizona, California, Idaho, Illinois, Mis-
souri, Nevada, Ohio, Oregon, Pennsylva-
nia, Texas, Utah, Virginia, Wisconsin. See
description of L. Hesperus for specific
localities of that species.

Hosts. — Diptera: Cecidomyiidae (Vig-
giani and Laudonia 1994). Lepidoptera:
Noctuidae, Pyralidae (Polaszek et al.
1998). The cecidomyiid record of larval
parasitism is a documented exception of
eggs serving as host in the Tricho-
grammatidae.

Lathromeris hesperus Pinto, new species

(Figs 50, 127, 231, 263)

Description. — Female. Body relatively
elongate, length 0.6-1.1 mm (N = 5), meta-
soma almost twice length of mesosoma.
Color dark brown to almost black except
petiolar area, vertex and front of head
yellow, antenna light brown, base and apex
of femora and tibiae and tarsomeres I and
II light yellow, fore wing disk colorless
with a broad fumate fascia extending
across wing behind MV and SV.

Antenna slender; length/width of scape,
pedicel and club: 38/11, 24/14, 68/14; Al
cupuliform, longer than disciform A2
which is closely appressed to CI. Club
narrow, surface with straight, obsolescent
longitudinal wrinkles; C1-C3 slightly wid-
er than long, C4 slightly longer than wide,
C5 much longer than wide, terminal pro-
cess short, 0.2 X length of C5, width of
segment at base of terminal process sub-
equal to width of process itself; C2-C5
relatively symmetrical, CI somewhat less
so; length/width of club segments: 14/18,
16/20, 19/20, 23/16, 43/8. Club sensilla
relatively sparse: CI and C2 with few APA,
APB, and 1 BPS near apex, C3 similar but
with BPS considerably below apex and
a few FS distally, C4 with FS, APB and BPS
also below apex, C5 apparently with 2 RS

Volume 15, Number 1, 2006

at apex attaining tip of terminal process; 2-
3 PLS on C4 and C5, terminal process with
a short UPP at apex. Mandible quadriden-
tate with anterior-most tooth markedly less
well sclerotized, lobiform. Maxillary palp
subcylindrical, c. 3x as long as wide, with
a narrow, cylindrical sensillum at apex (c.
0.4-0.5 X length of palp), terminal seta
slightly longer than palp.

Midlobe of mesoscutum with 2 pair of
moderately long, subequal setae, posterior
pair considerably more medially placed.
Scutellum with anterior pair of setae short,
only c. 0.2X length of posterior pair.
Sculpturing on midlobe consisting of rela-
tively large subpentagonal or hexagonal
cells except surface more longitudinally
rilled at middle of basal half; scutellum
with all sculpturing longitudinal. Propo-
deum relatively short, c. 2x as long as
metanotum at middle, discal area not
produced. Fore wing 2.5 X as long as wide,
venation attaining 0.6 FWL, fringe setae
moderately elongate, c. 0.3 X FWW; vena-
tion elongate, narrow, MV 2x as long as
PM, SV c. 0.3 X length of MV, slanting
away from MV at an angle slightly greater
:han 90 :', with only a slight constriction
between stigma and MV; MV with 3
elongate dorsal and 7 short ventral setae;
ipical half of PM and basal third of MV
poorly sclerotized, colorless; disk moder-
ately densely setose in area apical to
/enation, setae short especially posteriorly,
inear setal tracks present but poorly in-
iicated, linear tracks more distinct in
interior section of wing; costal cell without
setae. Hind wing with 3 linear setal tracks,
;etae short. Ovipositor moderately long,
lot extending beyond apex of metasoma,
3L/HTL = 2.0 (1.7-2.2) (N = 5), gonoplac
: 0.2 OL; hypopygium short, c. 0.2 OL.

Male. Body length 0.6 mm (N = l). As
emale except C5 without a terminal pro-
:ess and considerably shorter and wider
elative to C4 (subequal in length and 0.7X
ts width), C4 with APA and C5 with FS.
Metasoma with unique pair of lateral
ongitudinally arranged tubiform struc-

tures beneath surface of terga VI and VII,
each tube extending posteriorly from an
oval saclike structure near base of V to near
apex of VI, tubes then curving toward
midline but not obviously joining. GC with
length 0.5 X HTL, ADA c. 0.4 X GL, ventral
setae present, PAR absent; aedeagal AAP
present.

Types. — Holotype 9 and allotype S- UNITED
STATES. California: Riverside Co., Menifee
Valley (hills on W. side), 33 39' N, 117 13' W
(1800'); vi-7/13-1995 (holotype), vi-14/28-1995
(allotype); MT; J. D. Pinto, collr.; in NMNH.
Paratypes (2J, 99), same data except four with
additional dates (vii-12-1995, vii-31-1995); de-
posited in CNC, BMNH and UCRC.

Etymology. — Hesperus (Gr.), classical
name of the 'evening star', in reference to
the western distribution of this species.

Distribution. — Western United States;
Oregon, Nevada and Utah to southern
California

Comments. — Both sexes of Lathromeris
hesperus are readily distinguished from
congeners. In females C5 is very narrow
and narrows considerably towards the
apex where its width at the base of the
terminal process is subequal to the width
of the process itself. This results in the base
of the terminal process being difficult to
discern (Fig. 50). The short hypopygium,
the very dark brown coloration, fasciate
fore wings (Fig. 127), and the distinctive
mesosomal sculpturing also helps sepa-
rate females. Males are distinguished by
the pair of unique tubiform structures
beneath metasomal terga V and VI
(Fig. 263). Males of all other species known
to me either have large subquadrate
pustules or mamelliform evaginations as-
sociated with the apex of the metasoma
(Figs 262, 264). Females of Lathromeris
argentina, the only other described New
World species, has a considerably broader
C5, and lacks the fasciate fore wing. Its
male is unknown.

Material examined. — 549 (18 on slides), 7$ (6 on
slides). UNITED STATES. California: Del Loma,
Va mi. E (Erv Pigg property) (Trinity Co.); ix-1 /3-

100

Journal of Hymenoptera Research

1995; MT nr. forest edge; 29; S. Polly/G. Platner.
Granite Mountain Reserve (Granite Cove, c.
4200') (San Bernardino Co.); 34 48'N, 115
39'W; SP; 1<?; G. Platner. Lake Skinner, NE
(Riverside Co.); 33 35' 58" N, 117 01' 58" W (c.
1570'); iv-24/v-8/1997, vi-4/18-1996; MT in
'coastal sage scrub'; 39, 2 J; J. Pinto. Same except
33 36' 04" N, 117 02' 18" W (c. 1580'); v-22/vi-5-
1997, vi-18/vii-2-1998, vii-2/ 16-1998; 79, 2<J; J.
Pinto. Menifee Valley (hills on W. side) (River-
side Co.); 109, 36*; see Types [additional material
from type locality on cards: vi-7/ 13-1995, vi-14/
28, vi-28/vii-12-1995, vii-19/viii-l-1995; MT; 209;
J. Pinto]. Riverside (UCR campus); v-2-1984; 19;
J. LaSalle. Santa Rosa Plateau Ecological Reserve
(590 m) (Riverside, Co.), 33 32.524' N, 117
14.644' W; MT; vii-30/viii-14-2001; 19; PEET
Survey. Tamarack Valley (San Jacinto Mts.,
9120') (Riverside Co.); 33 48.61' N, 116 39.63'
W; SP; viii-13/ 17-2001; 19; J- Pinto/G. Platner.
Valyermo (Los Angeles Co.); viii-30-1989; SP;
19; R. Crandall. Nevada: Alamo, 17 mi. S
(Lincoln Co.); v-22-1995; 'scrn. swp. desert
veg.'; 19; J. Pinto. Winters, 11 km W (Cold
Canyon Reserve, Solano Co.); viii-1/ 15-1994;
MT, 'live oak woods'; 49; L. Kimsey. Oregon:
Brookings, near (Timeus Ranch, Curry Co.); 42
06' N, 124 17' W; ix-1/15-1995; MT; 10; M.
Wasbauer. Utah: Caineville, 6 mi. W (along
Fremont River, c. 4700') (Wayne Co.); vi-29-
1993; SP; 19; J- Pinto.

32. Lathromeroidea Girault

(Figs 6, 73-77, 144-146, 164-167, 180, 181,

183, 191, 196, 246-249, 261)

Lathromeroidea Girault 1912. Type species: La-
thromeroidea nigra Girault, by original desig-
nation.

Diagnosis. — Female. Body shape variable
but usually more elongate and slender than
similar genera (Uscanoidea and Burksiella).
Antenna with 2 anelli and a 5-segmented
club, CI shorter and narrower than C2 and
usually closely appressed. Fore wing 1.9-
2.5 X as long as wide, fore wing fringe 2:
0.20 X FWW in most species; venation
similar to that in Burksiella and Uscanoidea,
SV projecting apically beyond apex of MV;
disk moderately to densely setose, RSI,
usually distinct, typically straight, not
curving greatly from stigmal vein to in-

tersection with Cu tracks; propodeal disk
elongate, longer than metanotum, usually
at least twice as long.

Male. Eyes smaller than in female (cf.
Figs 165, 166). GC short, GL less that HTL,
without PAR, VS and aedeagal AAP,
ventral setae present.

Lathromeroidea is herein divided into
three groups (see Discussion).
Distribution. — Widespread.
Diversity. — Nine species. Seven have
been described; two new species, the only
named New World species, are added
here. Although exceptionally diverse in
the New World, none of its species has
been previously described. Doutt and
Viggiani (1968) did place odonatae in
Lathromeroidea, however it is herein re-
turned to Centrobiopsis (see above).

Discussion. — Lathromeroidea is similar to
and unsatisfactorily separated from Usca-
noidea. Differences are discussed under the
latter. Doutt and Viggiani (1968) suggested
alliance with Neolathromera as suggested by
similar wing and antennal structure. Nei-
ther Doutt and Viggiani nor I have seen
material of Neolathromera. In my opinion,
based on the original description, Neola-
thromera may be closer to Pten/gogramma.
Although the antennal club is 5 segmented
in all three, in Lathromeroidea CI is much
shorter than C2, not much longer as in
Neolathromera and Pterygogramma. Males of
Neolathromera are unknown.

As defined here, Lathromeroidea is a very
complex genus with high diversity. The
variation among species is considerable,
yet clear-cut divisions in the group are not
obvious. Rather than erect new genera,
perhaps prematurely, I divide the genus
into three informal phenetic species groups
A, B and C based on fore wing structure
alone. Groups A and B are widespread in
distribution; C is known only from the
New World. All previously described
species belong to Group A. Groups A and
C at least, appear to be associated with
aquatic habitats. Exemplar species of
groups B and C are described below.

olume 15, Number 1, 2006

101

Characteristics and comments regarding
ie three assemblages follow.

Group A. Fore wing (Fig. 144) c. 2.3 X as
mg as wide, fringe setae c. 0.3 X FWW;
enation attaining 0.5 FWL, with MV 1.5-
.8X length of PM, SV+MV/PM = 2.0-2.4,
pex of SC terminating above base of PM;
isk moderately densely setose, with a bas-
l convergence of setal track RSI and most
f the other tracks posterior to r-m; a line of
?tae present immediately anterior of the
?tinaculum; a narrow but distinct costal
sll present with more than 3 setae;
enation with a distinct constriction be-
veen the MV and stigma. Figs 73, 144.

This group includes the type species. All
2ven of the previously described species
f Lathromeroidea belong here. These spe-
ies occur in Australia, Europe and Asia,
he New World representatives of Group
. are very similar to those in other parts of
le world. Revisionary studies are needed
) determine which, if any, are conspecific
) described species.

Group B. Club segments usually more
symmetrical in female. Fore wing
ng. 145) often wider than in congeners,
.9-2.4 X as long as wide and fringe setae
ften shorter (0.2-0.4 X FWW); venation
ttaining 0.5 FWL, MV length 1. 5-2.0 X that
f PM; SV+MV/PM = 2.0-2.3, apex of SC
'rminating above base of PM; disk less
ensely setose than in the other groups,
dthout a conspicuous convergence of vein
acks basally, at most only RSI and Cu
'acks converging toward base of wing,
sually without a distinct line of setae
nterior to retinaculum, rarely more than 3
?tae in costal cell. Midlobe of mesoscutum
dth posterior setae slightly medial to
nterior pair; scutellum not abruptly de-
ressed anterolaterally; metasomal tergum

not longitudinally divided, its anterior
largin relatively straight. Figs 6, 74-76,
45, 164, 174, 181, 183, 191, 249.

This group includes the least modified
secies of the genus and the most likely to
e confused with Uscanoidea. It is consid-
rably diverse in the New World tropics

where perhaps it is the most frequently
collected group of Trichogrammatidae. A
single representative, L. exempilum, is de-
scribed below to characterize the group.
Although apparently not common else-
where, I have examined species belonging
here from North America, Australasia and
Africa.

Group C. Body much more heavily
sclerotized than congeners and other tri-
chogrammatids. Head of both sexes with
a unique deep and relatively large pit
immediately in front of anterior ocellus
(Figs 165-167). Fore wing (Fig. 146) nar-
rower, 2.3-2.5 X as long as wide, fringe
setae 0.3-0.4 X FWW; venation attaining 0.4
FWL, MV 1.3-1.6X length of PM, SV+MV/
PM = 1.5-1.8, SV sessile to MV or with only
slight constriction at base, SC and PM
confluent, PM closer to wing margin and
costal cell absent or extremely narrow; disk
densely setose, setal tracks indistinct or
obvious only in a narrow band anteriorly
and posteriorly, without a line of setae
anterior to retinaculum. Midlobe of mesos-
cutum with posterior pair of setae lateral to
anterior pair; scutellum strongly depressed
anterolaterally and appearing laterally
emarginate in dorsal view (Fig. 180). Meta-
somal tergum II (first visible) usually
elongate and longitudinally divided at mid-
dle, extending ventrally to overlap much of
anterior sternal plates, with a unique oval,
clear bulla anterolaterally (Fig. 196); last
sternum of both sexes divided apically with
each lateral lobe produced into an acute
posterior projection (Fig. 261). Figs 77, 146,
165-167, 180, 196, 246-248, 261.

The cephalic pit of Group C is unknown
in any other Trichogrammatidae. However
a very similar structure, termed the pre-
ocellar pit, occurs in the Scelionidae,
Megaspilidae and Ceraphronidae (Bin
and Dessart 1983). The ultrastructure of
this pit has been examined in the
scelionid Trissolcus bnsnlis (Woll.) by Isi-
doro and Bin (1994). In this species the pit
is manifested internally as an apodeme
connected to the protocerebrum by a bun-

102

Journal of Hymenoptera Research

die of microtubule-rich epidermal cells.
Isidoro and Bin assume that this structural
complex provides mechanical support for
the brain.

Group C is the most distinctive and is
known only from the New World. Its
differences suggest generic or subgeneric
status. Three traits alone, the strongly
sclerotized body, the preocellar pit and
the presence of metatergal bullae, are
unique within the family. Whereas the
limits between groups A and B, and
between Group B and Uscanoidea are not
clear, Group C would appear to be easily
characterized. Yet, as defined, Group A
includes certain species which have fore
wing structure typical of its group but
scutellar and certain metasomal character-
istics of Group C. Therefore, before parti-
tioning, I believe the entire Lathromeroidea
complex as well as its relationship to
Uscanoidea warrant careful study.

New World records. — Group(s) are indi-
cated after each record. Argentina: Mis-
iones (A/B). Belize (A/B). Bolivia (A/B).
Brazil: Goias (A/B), Rondonia (B). Canada:
Alberta (C), Ontario (C), Quebec (C).
Colombia (A, B). Costa Rica (A/B/C).
Ecuador (A/B). Guatemala (B). Mexico:
Chiapas (B), Michoacan (B), Tamaulipas
(B), Quintana Roo (A/B), Veracruz (A).
Panama (A/B/C). Paraguay (A). Peru (B).
United States: Arizona (A), Florida (A/B/
C), Georgia (A/C), Kentucky (C), Missouri
(A/C), New Mexico (A), South Carolina
(B/C), Utah (B). Venezuela (A/B/C). West
Indies (A/B). See descriptions of L. exem-
plum and L. gerriphaga for specific records
of those species.

Hosts. — Hemiptera: Gerridae (Henriquez
and Spence 1993, record for Group C).

Lathromeroidea exemplum Pinto,
new species

(Figs 145, 164, 174, 181, 183, 191)

This species is one of several belonging
to Section B. It was chosen for description
because it is the most commonly collected

species of the group. Body length and color
are based on two card-mounted specimens.
Description. — Body moderately slender,
length 0.4 mm. Color predominantly mod-
erately light brown. Female. Head broad,
0.6 as long as wide. Eyes relatively large,
occupying c. 0.8 distance from top of eye to
apex of clypeus; toruli placed between
eyes, c. half distance from top of head to
base of clypeus. Maxillary palp cylindrical,
almost 3x as long as wide, with an
elongate peg-like sensillum at apex c.
0.6 X length of both palp and terminal seta,
resp. Antenna with an elongate, narrow
club, club > 2x length of scape; measure-
ments (length /width) of scape, pedicel and
club: 50/8, 26/15, 110/18; scape narrow,
uniform its entire length; club outline
symmetrical, in shape of a long, narrow
cone. Club 5 segmented; CI very short,
transverse, closely appressed to C2, pri-
marily visible in medial view; club widest
at C2-C3; CI, C3 and C4 asymmetrical, C2
less so, C5 symmetrical, narrow, of rela-
tively equal width entire length except at
very apex; measurements (length/width)
of C1-C5: 4/15, 15/21, 30/20, 37/18, 46/
11. Club sensilla: BPS large, subglobose, at
or near apex of all segments; C2-C5 with 1,
1, 1, 4 PLS, resp., PLS on C2 strongly
curved; C2 and C3 with few APA; APB on
C2 (2), C3 (1) and near apex of C5 (1); C3-
C5 with several FS; C5 with 2 RS near apex
and setiform UPP at apex (usually curved);
longest FS c. 2.2 X maximum club width.
Mid lobe of mesoscutum and scutellum
each with 2 pair of stout, elongate setae;
posterior pair on midlobe more medial in
position than anterior pair. Propodeal disk
slightly produced posteriorly, 2x length of
metanotum; mesopleuron broad, without
pleural suture. Sculpturing of mesoscutum
and scutellum areolate, cells primarily
longer than wide and with elevated bor-
ders, most reticulae with a series of trans-
verse ridges within, ridges becoming lon-
gitudinal posteriorly on midlobe. Fore
wing 2.2 X as long as wide, venation
attaining c. 0.6 FWL, fringe setae c. 0.25 X

Volume 15, Number 1, 2006 103

FWW; MV elongate, narrow, straight, c. 1995, Malaise trap, #02.448, Alas Project".

1.7 X length of PM, both veins subequal in Paratypes deposited in BMNH, UCRC. All

width; PM diverging from MV forming types are on slides. The collector is not indicated

a distinct costal cell, base of PM posterior on any of the [YPes> but a11 are from collections

to apex of SC; SV arising from poster- of Project ALAS at La Selva.

oapical corner of MV, elongate, 0.4 x length Etymology.— Exemplum (L.) == example,

of MV, with a distinct, narrow neck Geographic distribution.— Known from

between stigma and MV; MV with 3 southern Mexico S to Peru,

elongate anterior and 4 shorter posterior Material examined.- 31o (29 on slides), 7; (all

setae on dorsal surface, with 6 ventral on slides). BELIZE. Toledo: Maya Mts., Bladen

setae; PM with 2 elongate setae and 2 Branch Forest Res. (150 m); viii-1/ 15-1989, ix-3-

campaniform sensilla at apex; disk moder- 1989; PT/MT; 49, 46*; M. Williamson. COSTA

ately densely setose with most setae asso- RICA. Alajuela: Reserva Rincon Forestal, Est.

ciated with linear setal tracks; RSI well Caribe (4°0 m), 10 53' N, 83 18' W; ii-19/20-

developed, slightly curved toward wing 2003; l<?'' J- Noyes. Heredia: La Selva; 79, 2; (see

base and converging on Cu tracks, tracks TyPes)- OTS"La Selva <100 m>; {A99l> YPT' l^ J-

anterior to Cu not extending to point of J^.1^".^ I *£ SW (5° m); [xM~

, , . , .,, - 1989, xu-1989/ii-1990; 29; P. Hanson. Guapiles,

convergence; basal track with 2 setae; 16 km W (480 m); x/xi-1989, xii-31 -1989; 29 1 ;;

costal cell with 1-2 setae; radial process at p Hanson Puutamws: Parque Nacional Cor.

base of PM obsolescent. Hind wing with covado (Estacion Sirena) (50 m); v/viii-1989; I9.

complete anterior and medial setal tracks, r. p. Golfo Dulce (Piedras Blancas, 24 km W)

posterior track consisting of a few basal (200 m); iv/v-1992, x/xi-1990; 39; P. Hanson,

setae only. Middle leg with an elongate, Same except: Piedras Blancas, 5 km W (100 m);

fine, tibial spur, spur length slightly greater xii-1990; 19. ECUADOR. Napo: Sucumbios Riv-

than that of first tarsomere. Ovipositor er> Sacha Lodge, 0 30' S, 76 30' W; vi-18/23-

relatively long, OL/HTL = 1.78 (1.5-2.0); 1994' MT- l?> p- Hibbs- GUATEMALA. Suchite-

gonoplac comprising c % OL pequez: Suchitepeque, Finca Moca Grande; ii-23/

Male. Eyes considerably smaller than in ™: lft Dv Q*^ MEXICO. Q,„„(„„„

c , . „ , „J„ ,. , , Roo: Nuevo X-can, 10 km S; xn-6-1993; swp.

female, occupying 0.6-0.7 distance from ,. ., . , ., 1rt , , . ' T\

, rj ° . , trail in secondary forest ; I9; L. Masner. PERU,

top of eye to apex of clypeus. Antennal LoretQ. Teniente Lopez (220 m)# 20 36% 76

club shorter than in female, length 1.5X 07-W; vii.22-1993; 29; R. Leschen. VENE-

length of scape, scape more tumid, mea- ZUELA. Aragua: Cumboto (cacao plantation)

surements (length/width) of scape, pedicel (50 m); v-13-1999; MT; 49; J. Garcia / R.

and club: 47/14, 30/18, 68/19; C5 shorter Montilla.

relative to other segments than in female, Comments. — Section B of Lathromeroidea

subequal to C4 in length. Club sensilla includes at least 15-20 undescribed species

similar to female except RS absent on C5 in the New World. L. exemplum females are

and fewer FS and APA. GC simple, separated from all that 1 am aware of by

rounded at base, narrowing at middle, the following traits: antenna elongate and

ADA 0.4 X GL, with ventral spines posi- narrow, with large, subglobose BPS; sculp-

tioned c. half distance from base to apex; turing of midlobe of mesoscutum consist-

GL/HTL = 0.6. ing of relatively large cells, usually slightly

_ Tr , , ^^o^a n,^4 r, j- t longer than wide, with transverse ridges

Types. Holotype 9. COSTA RICA. Heredia: La .f\ ir. A' , ?.

Selva; vii-17-1995; M08.405; in INBC. 59 para- Wlthm (Fl§- 183); ovlPosJtor moderately

types also from La Selva as follows: 19, same lon§ (OL/HTL 1-5-2.0X) extending only

data as holotype except 'M04.402'; lo, same data slightly beyond the apex of the metasoma.

as holotype except without date and with label Sculpture pattern is a convenient feature

indicating 'OTS-La Selva, M05.403'; 29, as for identifying slide-mounted specimens of

holotype except 'OTS-La Selva, 100 m, ix-14- L. exemplum. The few other known species

204 Journal of Hymenoptera Research

with a similar pattern are separated by BPS ly in front of anterior ocellus, diameter of

shape (narrow, subfusiform) or a much pit c. 1/6 that of ocellus,

longer ovipositor (> 3x HTL). The species Female. Eyes large, occupying more than

with the longer ovipositor also has a pro- 0.90 distance from top of eye to apex of

portionately longer gonoplac. In L. exem- clypeus (Fig. 165). Antenna relatively short,

plum the gonoplac is a fourth the length of with length /width of scape, pedicel and

the entire ovipositor; in the other species it club: 70/16, 35/17, 67/23; scape widest at

is a little over half its length. I am not basal half, tapering to c. half maximum

convinced I can separate males of L. width at apex; pedicel relatively elongate,

exemplum from those of the latter species, anellar segments distinct, Al slightly wider

The description of the male in the species than A2; club widest at C3, subfusiform,

description is based on two specimens segments all distinct and only slightly

belonging to the same series as the female asymmetrical, all segments except subco-

types. The Material examined section only nical C5 at least slightly wider than long;

includes males collected with identifiable length/ width of C1-C5: 11/15, 20/21, 14/

females. 24, 20/22, 20/18. Club sensilla: relatively

elongate APA and a few FS on all

Lathromeroidea gerriphaga Pinto, segments; C2-C5 with 1, 1, 2, 4 PLS, resp.;

new species C5 with 2 RS near UPP not visible;

(Figs 77, 146, 165, 166, 180, 196, 248, 261) BCp ^ apex rf gegmFents narroW/ subda.

As indicated below there are at least vate. Midlobe of mesoscutum and scutel-
three species in Section C of Lathromeroidea. lum each with two pair of elongate,
The species described was chosen be- relatively fine setae. Propodeal disk slight-
cause it has been the subject of a detail- ly produced and inclined posteriorly,
fed behavioral study by Henriquez length c. twice that of metanotum. Fore
and Spence (1993). In that paper, L. wing 2.5X as long as wide, venation
gerriphaga was referred to as Lathromeroidea attaining 0.4-0.5 FWL, fringe setae c. 0.3 X
sp. nov. FWW; venation straight, both PM and MV

Description. — Body moderately elongate, immediately adjacent to anterior margin of

length 0.7-0.8 mm. Color primarily dark wing, only SC placed slightly behind

brown with antenna and lateral areas of margin; SC and PM confluent, PM and

mesosoma distinctly lighter brown, legs MV slightly disjunct; MV widening grad-

primarily yellow to yellow brown except ually and slightly from base to apex, c.

fore and hind coxae light brown, middle 1.2X length of PM; PM subequal in width

legs entirely yellowish. Surface shiny, very to basal portion of MV; SV present but

heavily sclerotized throughout, with the relatively short, subsessile to MV, with

following areas on head and mesosoma only a slight constriction at base of stigma,

distinctly alveolate: head posteriorly and its length subequal to apical width of MV;

below anterior ocellus, pronotum, lateral MV with three elongate anterior and c. 6

areas of mesosoma including axillar and shorter posterior setae dorsally, and 6-7

axillular surfaces, propodeum, and coxae; setae ventrally; PM with two dorsal setae

these areas contrast with the relatively and two campaniform sensilla at apex; all

alutaceous mesoscutum and scutellum; venation setae fine; disk densely setose

metanotum somewhat transitional be- with distinct linear setal tracks obvious

tween the smoother scutellum and alveo- only in relatively narrow anterior and

late propodeum; metasoma smooth, shiny, posterior section, remainder of wing with

except metasomal tergum II weakly alveo- exceptionally dense setation, area posterior

late anteriorly. Head excavated between to venation without setation except three in

eyes with a deep preocellar pit immediate- basal vein track and one or two additional

Volume 15, Number 1, 2006

105

setae between retinaculum and venation.
Hindwing narrow, anterior and middle
setal tracks complete, a posterior track
present only apically. Middle leg with an
elongate fine, tibial spur, its length slightly
greater than that of first tarsomere. Meta-
somal tergum II elongate, almost as long as
remaining segments combined, with a clear
oval bulla in anterolateral corners. Ovipos-
itor moderately long, OL/HTL = 0.8-0.9
(N = 3).

Male. Eyes much smaller than in female,
occupying less than 0.7 the distance from
top of eye to apex of clypeus (cf. Figs 165,
166). Antenna without RS on C5 and only 1
and 2 PLS on C4 and C5, resp. GC (Fig. 248)
with length 0.4 X HTL (N = 2), c. twice as
wide at base as apically, rather abruptly
narrowing at basal 0.4, basal margin trans-
verse and distinctly flanged laterally; ADA
extending c. half length of capsule; a pair of
stout ventral setae positioned 0.6 distance
from base to apex of GC.

Types.— Holotype 9, allotype > CANADA.
Alberta: nr. Dunstable, Kirchner's Pond; ix-4-
1990; ex Limnoporus dissortis Drake and Harris
(Gerridae) eggs; N. H. Moreno, collr.; deposited
in CNC. Thirteen paratypes (10o, 3 j) with same
data, deposited in CNC, UCRC.

The type material of L. gerriphaga comes
from collections used by Henriquez and
Spence (1993) for their study. In that paper
they provide greater precision for the type
locality than indicated on the collection
labels: "Kirchner's East Pond, located c.
100 km NW (114 06'W, 53 57'N) of
Edmonton, Alberta".

Etymology. — The name gerriphaga is
based on Gerridae eggs being the recorded
host of the species.

Geographic distribution. Alberta, Canada
and southeastern United States.

Materia! examined. — 159, 4 o* (all except 59 on
slides). CANADA. Alberta: nr. Dunstable; 11 9,
4^ (see Types). UNITED STATES. Florida: Green
Swamp (Lake Co.); vii-30-1987; MT; 19; V.
Gupta. Archbold Biological Station (Highlands
Co.); vii-15/ix-l-1987; I9; D. Wahl. Georgia:

Sapelo Island, Saranna; vi-28/v-19-1987; FIT;
19; BRC Hym. Team. South Carolina: Clemson,
Cherry's Crossing (Pickens Co.); ix-6/ 13-1987;
lo*; J- Johnson.

Comments. — There are at least two addi-
tional species in Group C of Lathromeroidea.
Another species from the United States
(southeastern states) and Canada (Ontario
and Quebec) is similar to L. gerriphaga but
can be distinguished by color. In that
species the mesosoma is dark brown
throughout, not distinctly lighter laterally.
Also its legs are brown, not yellowish as in
L. gerriphaga. The fore wing provides an
additional difference. In the undescribed
species the premarginal vein is slightly
shorter and placed slightly posterior to the
anterior wing margin forming a narrow
and setose costal cell. In L. gerriphaga the
premarginal vein coincides with the ante-
rior wing margin its entire length. A third
species occurs in Central America and
northern South America. This species is
easily distinguished from both of the
above. Its fore wing disk is densely setose
throughout (setal tracks not distinct any-
where on disk), its hindwing is also
densely setose and without obvious linear
setal tracks, and the stigmal vein is much
shorter and sessile to the marginal vein,
similar to the condition in Aphelinoidea. In
addition, the male genitalia of this species
are attenuate and narrowest at its very
base, whereas in both North American
species the genital capsule is broadest at
the base (as in Fig. 248).

33. Monorthochaeta Blood

Monorthochaeta Blood 1923. Type species: Mono-
rthochaeta nigra Blood, by monotypy. Blood
and Kryger 1928 (redescription).

Diagnosis. — Female. Antenna with 2 an-
elli, 2 funicular and 3 club segments;
funicular segments subequal in length
and width, closely appressed; club rela-
tively narrow, elongate, slightly narrower
than funicle; all postanellar segments ex-
cept Fl with at least 1 PLS. Fore wing c.

106

Journal of Hymenoptera Research

2.4 X as long as wide; venation with well
developed SV, a radial process present;
disk densely setose, linear setal tracks not
clearly distinguished from surrounding
setae, RSI absent. Legs with first tarsomere
longer than others.

Male. Wingless (M. nigra) or winged as in
female (M. galatica Nowicki). Antenna club
2 (M. nigra) or 3 segmented (M. galatica). GC
(M. nigra) with PAR absent, VS present and
asymmetrical; apodemes of AAD present.

Distribution. — Primarily Palearctic. A
single specimen is known from the Nearc-
tic (Canada).

Diversity. — Two species of Monortho-
chaeta, M. niger and M. galatica Nowicki, are
known from the Palearctic (Nowicki 1940).
The unique specimen from the New World,
a female, cannot be separated from M. niger.
Without males, however, its identity re-
mains questionable. The South American
species (platensis De Santis) assigned to
Monorthochaeta by Doutt and Viggiani
(1968) is now placed in Burksiella (see above).
Discussion. — In the New World, Mono-
rthochaeta is most likely confused with
Ufens. Females of both have similar anten-
nal structure but are separated by the fore
wing. In Ufens the wing is considerably
broader and has distinct linear setal tracks
including an RSI. Males of Ufens are
further separated by the four-segmented
club. The separation of Monorthochaeta and
Densufens, described from China, requires
further study. Lin (1994) placed Densufens
close to Monorthochaeta but separated it by
the three-segmented club in males. Al-
though the club is two segmented in males
of M. niger, it also is three segmented in M.
galatica (Nowicki 1940). I have not exam-
ined M. galatica and have no opinion as to
whether it is congeneric with the type
species of Densufens. However, it is clear
that the two nominal genera cannot be
separated as currently defined.

The single New World record of Mono-
rthochaeta was discovered too late to allow
figures. However, males and females of the
genus are well illustrated in the literature

(Nowicki 1940, Doutt and Viggiani 1968,
Viggiani 1971).

New World record. — Canada: Ontario
(Ancaster).

Hosts. — Coleoptera: Chrysomelidae (La-
beyrie 1962).

34. Nicolavespa Pinto

(Figs 11, 51, 128, 232, 233)

Nicolavespa Pinto 2005. Type species: Nicolavespa
theresae Pinto, by original designation.

Diagnosis. — Female. Antenna with 2 anelli,
2 funicular and 3 club segments; Fl short,
transverse, closely appressed to F2; F2 more
elongate, subcylindrical; club with a distinct
terminal process. Fore tibia without spines
on dorsal surface. Fore wing moderately
broad, usually slightly less than twice as
long as wide, venation with SV well de-
fined; disk moderately densely setose with
distinct setal tracks including RSI.

Male. Antennal club without an apical
process. GC with VS, ventral setae and
aedeagal AAP present, PAR apparently
absent.

Distribution. — United States S to Central
America.

Diversity. — Nicolavespa contains two spe-
cies, the widespread N. theresae and N.
luiseno Pinto, restricted to southern Cali-
fornia.

Discussion. — Nicolavespa is the only
known genus of Trichogrammatidae hav-
ing a two-segmented funicle and a female
club with a distinct terminal process.
Superficially Nicolavespa resembles Chaetos-
tricha. Body shape is similar in both as is
antennal segmentation and general struc-
ture of the fore wing (Pinto 2005). Unlike
Nicolavespa, however, Chaetostricha lacks
a terminal process, the fore tibia has
a prominent thornlike spine on its dorsal
surface, and its genitalia lack aedeagal
apodemes as well as volsellae. Nicolavespa
may be more closely related to Lathromeris.
Although the antenna of Lathromeris lacks
a funicle and has a five segmented club,
a terminal process is present in females.

Volume 15, Number 1, 2006

107

Also, as in Nicolavespa, the male genitalia
have aedeagal apodemes. Other details of
the genitalia are not similar however, in
that parameres but apparently not volsel-
lae occur in Lathromeris.

Records. — Costa Rica. Guatemala. Mex-
ico: Chiapas, Guerrero, Jalisco, Morelos,
Sinaloa. United States. Arizona, California,
Georgia, Illinois, Maryland, Oklahoma,
South Carolina, Texas. West Indies. See
Pinto (2005) for detailed collection records.

35. Pintoa Viggiani
(Figs 52, 129, 184, 185)

Pintoa Viggiani 1989. Type species: Pintoa
nearctica Viggiani, by original designation.

Diagnosis. — Female. Antenna elongate,
with 1 anellus, 1 funicular and 3 club
segments; CI partially fused to C2, Fl with
a strongly curved J-shaped PLS; BPS
narrow, apically attenuate. Eyes reddish.
Midlobe of mesoscutum with 1 or 2 pair of
elongate setae; scutellum with a lateral and
medial pair of setae, lateral pair small to
minute. Fore wing narrow, 3.0-3.5 X as
long as wide, fringe elongate >0.5X FWW;
venation elongate, attaining 0.6 FWL, SV
distinct; disk moderately densely setose,
linear setal tracks present but not always
distinct, RSI absent.

Male. Antenna more elongate than in
female, CI not partially fused to C2; club
segments each with a basal whorl of
extremely elongate, erect FS. GC short,
tubular, enlarged basally, with a relatively
large ADA, VS and PAR absent, ventral
setae present; aedeagal AAP present.

Distribution. — Known only from the
New World (Canada to Argentina).

Diversity. — One species, P. nearctica, is
described. Its distribution spans that of the
genus. A second, undescribed, species
occurs from Florida to Ecuador. It is known
from females only.

Discussion. — Pintoa is not clearly related
to any other trichogrammatid genus.
Greatest phenetic resemblance is with
Oligosita. In both, the body is relatively

slender, the antenna has a single funicle
and a three-segmented club, the fore wing
is narrow with straight, elongate venation
and it lacks distinct linear setal tracks.
Pintoa is separated from Oligosita and all
other Oligositini by the reddish eyes (black
in oligositines), the presence of the J-
shaped PLS on the funicle, two pair of
setae on the scutellum (one pair in oligo-
sitines), and male genitalia (ADA much
reduced and apodemes absent in oligosi-
tines). The type species is well illustrated
by Viggiani (1989).

Among other differences, the two pair of
elongate setae on the mesoscutal midlobe
will distinguish the undescribed species of
Pintoa from P. nearctica which has only
a single pair.

Records. — Argentina: La Rioja. Belize.
Canada: Ontario, Quebec. Colombia. Cos-
ta Rica. Ecuador. Guatemala. Mexico:
Morelos, Nuevo Leon, Sinaloa Tamaulipas,
Quintana Roo. United States: Arizona,
California, Florida, Georgia, Louisiana,
Maryland, Missouri, North Carolina, South
Carolina, Oklahoma, South Carolina, Tex-
as, Virginia. West Indies.

Hosts. — Unknown.

36. Pseuduscana Pinto, new genus

(Figs 53, 54, 130, 234, 235)

Type species. — Pseuduscana sola Pinto, n. sp.

Description. — Small, length between 0.3-
0.4 mm; color brown with varying degrees
of white on mesosoma. Female. Antenna
with 1 anellus and 4 club segments. Club
subcorneal, narrow and elongate; segments
relatively symmetrical; CI short, transverse,
more than twice as wide as long, only c. 1 /3
length of C2; C2 about as long as wide, not
tapering to apex; C3 subequal in length to
C2, tapering slightly to apex; C4 much
narrower, c. twice as long as wide; C1-C3
of similar width, C2-C4 of similar length; all
club segments with conspicuous, large,
subglobose BPS at apex; C2 with a ring of
elongate FS at base; C3 with a basal and an
apical ring of FS. Maxillary palp 1 segment-
ed. Midlobe of mesoscutum usually with

108

Journal of Hymenoptera Research

only 1 pair of setae; scutellum with 2 pair
but anterior pair shorter than posterior pair.
Fore wing as in Uscana except narrower and
with longer fringe setae (length 0.2-0.3 X
FWW), also less densely setose with setal
tracks including RSI more conspicuous.
Hind wing with only an anterior and medial
setal track. Ovipositor short, shorter than
hind tibia; hypopygium present, of varying
length.

Male. Antenna with only a single ring of
FS on C3. GC with ADG completely
consolidated into GC but otherwise ex-
ceedingly variable (shape of GC, size of
ADA, presence or absence of PAR and VS,
and their expression when present vary);
ventral setae absent.

Etymology. — Name based on similarity
of the new genus to Uscana; gender
feminine.

Distribution. — Worldwide. In the New
World, from Canada to Argentina.

Diversity. — Based on male genitalia, I am
able to discern about 20 species of Pseudus-
cana in the New World; two of these occur in
the United States. Only one species, P. sola,
is described below. The greatest genitalic
and presumably species diversity occurs in
the New World. The few extralimital males
examined do not differ from one another
appreciably; all have simple genitalia with-
out volsellae and parameres.

Discussion. — Pseuduscana can only be
confused with Uscana and Brachista from
which it differs primarily in details of the
antennal club. In Pseuduscana CI is much
wider than long and considerably shorter
than C2, CI also lacks PLS, and all club
segments have large conspicuous BCP
sensilla. In Uscana and Brachista, CI is as
long or longer than C2 and, in females at
least, always bears at least one PLS.
Although BPS sensilla are present in these
two genera they are considerably smaller
and less conspicuous (cf. Figs 53, 61). In
addition, most Uscana and Brachista are
almost uniformly brown in color; most
Pseuduscana are distinctly bicolored, brown
with a considerable amount of white on the

mesosoma including much of the legs.
Certain Chinese species of Uscana have
relatively narrow fore wings with longer
than typical fringe setae and in this way
resemble Pseuduscana. Based on their de-
scriptions, however, characteristics of CI
are as in other Uscana (Lin 1994). Color
pattern and general body shape are similar
in some species of Haeckeliania but the
latter is separated by numerous features
including its five-segmented club and two-
segmented maxillary palp.

Pseuduscana is distributed worldwide but
greatest known diversity is in the New
World. External morphology of all species is
exceptionally homogeneous; yet the male
genitalia are more variable than in any other
genus of Trichogrammatidae that I am
aware of except perhaps Ufens. In some,
the genitalia are simple, consisting of
nothing more than a simple tubular capsule
(as in Figs 242, 243). In most, however,
parameres and volsellae are present (as in
Fig. 235) and their conformation takes on
a large number of possibilities. The shape of
the capsule and size of the ADA also vary
considerably. As in Ufens and Trichogramma
the taxonomy of this genus will depend
almost entirely on males.

New World records. — Argentina: Chaco,
Formosa, Misiones, Salta. Belize. Bolivia.
Brazil: Mato Grosso, Rondonia, Sao Paulo.
Canada: British Columbia, Ontario, Que-
bec. Colombia. Costa Rica. Ecuador. Gua-
temala. Mexico: Chiapas, Quintana Roo,
Tamaulipas. United States: California,
Florida, Georgia, Maryland, Missouri,
North Carolina, Oklahoma, South Caro-
lina, Texas, Virginia. Venezuela. West
Indies. See description of P. sola for specific
localities of that species.

Pseuduscana sola Pinto, new species

Description. — Small, body length 0.35-
0.40 mm. Color distinctly bicolored, with
metasoma, pronotum and hind legs brown,
mesosoma including fore and middle legs
white, head and antennae light brown.

Volume 15, Number 1, 2006

109

Male. Mandible tridentate with 2 strong
sickle-shaped posterior teeth and 1 shorter,
more blunt anterior tooth. Maxillary palp
subcylindrical, c. 2X as long as wide, apex
with terminal seta c. 2X length of segment,
and a narrow, digitiform sensillum c. 0.4
segment length. Antenna with length/
width of scape, pedicel and club: 32/10,
22/13, 57/13; club narrow, subfusiform
with C2 slightly wider than CI and C3,
tapering evenly to apex; measurements
(length/width) of club segments (C1-C4):
6/11, 20/14, 21/12, 20/8, C2-C4 moder-
ately asymmetrical. Club sensilla: CI -
APB, 1 BPS, 1-2 APA; C2 - 1 PLS, 3 BPS,
2 FS; C3 - 1 PLS, 1 BPS, 4 FS; C4 - 2 PLS, 1
BPS, 1 FS, with PLS extending slightly
beyond apex of club.

Midlobe of mesoscutum with 1 pair of
elongate setae laterally at anterior fifth.
Scutellum with 2 pair of setae, anterior pair
c. half length of posterior pair, posterior
pair shorter and less robust than mesoscu-
tal setae; sculpturing on both structures
consisting of relatively large cells without
obvious corrugations within, cells variable
in shape, most longer than wide. Propodial
disk not produced posteriorly, subequal in
length to metanotum. Fore wing 2.1 X as
long as wide, venation attaining 0.5 FWL,
fringe setae elongate, 0.5 X FWW; MV
elongate, straight, 1.5 X length of PM, both
veins subequal in width; PM diverging
only slightly from MV, with 2 setae and 2
campaniform sensilla at apex; SV elongate,
c. half length of MV, arising from poster-
oapical corner of MV, with a distinct
constriction basal to stigma; MV with 3
elongate setae anteriorly and 1-3 setae
posteriorly (only 1 in holotype); disk
moderately densely setose, with almost
all setae associated with linear tracks; RSI
present, straight; basal track with 2 setae;
usually only 1 seta at apex of narrow costal
cell; radial process present but weakly
indicated. GC relatively large, GL 0.7-0.8
HTL, dorsal outline of genital capsule
somewhat paddle shaped, 0.3 X as wide
as long, narrowest aspect at apex c. 0.3 X

greatest width; entire apex of genitalia
strongly curved ventrally; ADA large,
occupying 0.65 X length and 0.75 X width
of capsule; PAR and VS well developed; VS
elongate, digitiform, adjacent, arising in
basal half of GC (c. 0.4 distance from base),
their length c. half GL, asymmetrically
tapering to a point at apex, reaching apical
0.9 of GC; PAR shorter, also attenuate at
apex, attaining apical 0.8 of GC.

Female. As male except antennal club with
more and longer FS, 2 whorls of FS on C3,
longest FS 2.8 X (N = 2) greatest antennal
width. Ovipositor very short, confined to
apex of metasoma, OL/HTL = 0.60 (N=2).

Type.— Holotype J: UNITED STATES. Mary-
land: Port Republic (Calvert Co.); viii/ix-1986;
FIT; Sharkey /Munroe; in CNC.

Etymology. — Sola (L.), sole or only.

Geographic distribution. — Southeastern
United States and Argentina.

Material examined. — 39, 46* (all on slides).
ARGENTINA. Misiones: Parque Nac. Iguazu,
Pto. Canoas (200 m); xii-8-1990/1-6-1991; 'hill
forest'; lo, 1& S./J. Peck. UNITED STATES.
Georgia: Calhoun (Gordon Co.) (285 m); 34 29'
09" N, 84 54' 22" W; v-16-2002; 16*; D. Yanega.
Sapelo Island; vi-29/vii-18; 1987; 'live oak
forest'; 1^; BRC Hym. Team. Maryland: Port
Republic; 1 j (see Type). Prince Frederick, 7 km
S; v-7/vii-7-1987; FIT, 'hardwood forest'; 2<?;
BRC Hym. Team. Texas: College Station, Lick
Creek Park; x-1 1/18-1987; MT; 1 y, R. Wharton.

Comments. — As indicated, Pseuduscana is
a large but homogeneous group. Although
diversity of male genitalia is extensive,
other characters vary minimally. At least
one other form, occurring in Argentina
(Misiones) and Mexico (Chiapas), has male
genitalia similar to P. sola. These may be
conspecific but are not treated as such
because of minor differences in sculpturing.
In P. sola the mesoscutal and scutellar
reticulae are relatively smooth within; in
the similar form, the reticulae are longitu-
dinally and transversely corrugated within.

Because of considerable morphological
homogeneity except for male genitalia,
females of Pseuduscana are not easilv

Journal of Hymenoptera Research

distinguished. In this respect the genus
very much resembles Trichogramma and
Ufens, two other genera with exceptionally
diverse male genitalia. Females included in
the Material examined section either were
collected with males and/or have no
obvious structural differences from males
of P. sola. Unfortunately, most of the
specimens collected of this genus are
females. Based on the relatively few males
available only two species can be recog-
nized in the United States, P. sola and an
undescribed species with very simple
genitalia lacking both volsellae and para-
meres. The latter also occurs in the South-
east and may be sympatric with P. sola. The
above description of females is based on
two specimens from near Prince Frederick,
MD, close to the type locality.

37. Pteranomalogramma Viggiani and
Velasquez

Pteranomalogramma Viggiani and Velasquez,
2004. Type species: Pteranomalogramma singu-
lare Viggiani and Velasquez, by original
designation.

Diagnosis. — Female. Antenna with 2 an-
elli, 2 funicular and 3 club segments;
funicle subglobose, Fl very short, closely
appressed to F2, F2 with PLS. Maxillary
palp 1 segmented. Midlobe of mesoscutum
and scutellum each with 2 pair of very
short setae. Fore tibia with dorsal margin
bearing 4-5 spines at apex; all tibiae robust.
Fore wing unique, almond shaped, apical
margin pointed; disk with area behind
venation fumate and without setae, setae
arranged in linear tracks in area apical to
venation, RSI absent; fringe setae elongate,
length >0.5X FWW; venation with SV
distinct, constricted between stigma and
MV, slanted diagonally away from MV.
Ovipositor very short.

Male. — Unknown.

Distribution. — Known only from Vene-
zuela.

Diversity. — Monotypic.

cussion. — Pteranomalogiwnma is known
5 single female. The almond-shaped

fore wing coupled with antennal structure
separates it from all other New World
genera. The spinose fore tibia also is
unique. The fore tibia of certain genera
such as Chaetostricha and Zaga also are
spinose but in these taxa the spines are
placed at the middle or along the entire
dorsal surface (Figs 176-178), not clus-
tered apically as in Pteranomalogramma.
Because males are unknown, placement of
this genus in the Chaetostrichini is based
solely on the antennal structure (first
postanellar segment very short, closely
appressed to second) which characterizes
several genera in this tribe.

I became aware of Pteranomalogramma as
this work was nearing completion and have
not had the opportunity to examine the type
species. Its treatment is based solely on the
original description and associated figures
(Viggiani and Velasquez 2004).

Records. — Venezuela (Aragua: Choroni).

Hosts. — Unknown.

38. Pten/gogramma Perkins

(Figs 55, 94-96, 131)

Pten/gogramma Perkins, 1906. Type species:
Pten/gogramma acuminata Perkins, by mono-
typy.

Abbelloides Brethes 1928. Type species: Abbel-
loides marquesi Brethes, by monotypy. Doutt
and Viggiani 1968. De Santis (1970, as
synonym).

Diagnosis. — Female. Antenna with 2 anelli
and 5 club segments; CI longer than C2; C2
transverse, shortest segment of club, only
club segment without a PLS. Propodeal disk
subtriangular, distinctly longer than meta-
notum. Fore wing 1.9-2.3X as long as wide,
fringe setae of variable length; venation in
most New World species with MV evenly
curved at apex to form SV; disk sparsely to
moderately densely setose, with most setae
arranged in linear tracks, RSI present,
membrane immediately apical to RSI usu-
ally subglabrous, area posterior to MV with
foliate sensilla ventrally. Hind wing disk
usually with only a middle row of setae.

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111

Male. GC with a pair of very short,
bilobed structures lateroventrally near
apex (parameres?), ventral setae present;
aedeagal AAP present.

Distribution. — Australia, Asia and in the
New World, the West Indies, Mexico, and
South America.

Diversity. — Pterygogramma includes 12
species. The genus is primarily Australa-
sian. Three species are described from
the New World: P. marquesi (Brethes)
from Brazil, P. pallidipes (Girault) from the
West Indies, and P. membraciphagum Vig-
giani from Argentina and Brazil. The types
of all three were examined. A few unde-
scribed species are in collections. The
genus is uncommonly collected in the
New World.

Discussion. — Pterygogramma appears to
be most closely related to the Australian
genera Thoreauia and Paruscanoidea. The
Japanese genus Neolathromera, although
virtually unknown, may also be related
based on its antennal structure. In the New
World however, Pterygogramma is most
likely confused with Latliromeroidea. Body
structure is similar in both, wings are of
similar dimension, and they share a five-
segmented club. Unlike Pterygogramma,
however, C2 is considerably longer than
CI in Latliromeroidea, not much shorter.

Most New World species of Pterygo-
gramma differ from congeners in venation
structure. In Australasian species the mar-
ginal vein ends abruptly rather than curv-
ing onto the stigmal vein. The South
American species P. marquesi is unique from
all other species in that the premarginal and
marginal veins are confluent, the midlobe of
the mesoscutum has three pair of setae, and
the scutellum is extremely setose with six
pair of setae (Fig. 95). I am unaware of any
other trichogrammatid with more than two
pair of scutellar setae. P. marquesi also
differs from most congeners in having
a more setose hind wing; its male genitalia,
however, are typical for the genus (Fig. 96).

New World records. — Argentina: Salta.
Brazil: Rio de Janeiro, Santa Catarina.

Bolivia. Ecuador. Guatemala. Panama.
Mexico: Quintana Roo. West Indies.

Hosts. — Hemiptera: Aetalionidae (Brethes
1928), Cicadellidae (Perkins 1906).

39. Tumidiclava Girault

(Figs 10, 56, 57, 132, 236, 237)

Tumidiclava Girault 1911. Type species: Tumidi-
clava pulchrinotum Girault, by original desig-
nation.

Orthoueura Blood 1923. Type species: Orthoiicum
bimaculata Blood, by monotypy. Blood and
Kryger 1928 (redescription).

Orthoneurella Blood and Kryger 1929 (;/. n. for
Orthoneura Blood 1923, nee Macquart 1829).

Diagnosis. — Female. Antenna with 2 anelli
and a suboval club of 2-3 segments; club
with an elongate terminal process at apex.
Fore wing moderately wide, slightly more
than twice as long as wide; venation
similar to Aphelinoidea, relatively short
(extending to c. 0.4 FWL) with stigma
broad and very short, sessile to MV, MV
with only 3 anterior setae on dorsal
surface, posterior setae absent; disk usually
moderately densely setose, R, RS2 and Cu2
usually distinguishable, remainder of sur-
face with setae scattered and not arranged
in linear tracks. Hind wing with complete
anterior and middle setal tracks, posterior
track usually absent or incomplete. Hypo-
pygium well developed, with a relatively
narrow posterior prolongation.

Male. Antennal club with 2-5 segments,
terminal process absent. Metasomal ter-
gum VII with a pair of lateral suboval or
reniform pustules posterior to spiracles,
tergum VI with or without similar struc-
tures. GC truncate or emarginate basally,
PAR present but usually reduced, VS
absent; aedeagal AAP present.

Distribution. — Worldwide. Known to oc-
cur from Canada to Argentina in the New
World. Uncommonly collected in the tropics.

Diversity. — There are 18 species of Tumi-
diclava described. Only two occur in the
New World. T. pulchrinotum is broadly
distributed in North America, and T.

112

Journal of Hymenoptera Research

pampeana De Santis occurs in Argentina.
There are a modest number of undescribed
species.

Discussion. — Tumidiclava is superficially
similar to Aphelinoidea. Characteristics of
the fore wing, antenna and male genitalia
are similar. In both, a funicle is absent, the
fore wing is only moderately wide, the
stigma is sessile to the marginal vein and
the fore wing discal setation is relatively
dense and not uniformly arranged in
linear tracks. The presence of the terminal
process on the club of Tumidiclava
females and the pustulate metasomal ter-
gum VII in Tumidiclava males separate
these genera. They also are distinguished
by differences of the fore and hind wings
(Pinto 1997a).

It is likely that Tumidiclava is more
closely related to Lathromeris. Although
fore wing structure does not suggest this,
females of both have a terminal process on
the antennal club, and the males have
similar metasomal modifications (as in
Fig. 264). Male genitalia and hypopygial
structure are similar in both groups also
(see Lathromeris).

Although most Tumidiclava are charac-
terized by the moderately densely setose
fore wing, I have examined undescribed
forms with relatively sparse discal setae; in
one species from Turkmenistan the disk is
almost glabrous.

New World records. — Argentina: For-
mosa, Jujuy, La Pampa, La Rioja, Salta.
Belize. Canada: Alberta, British Columbia,
Prince Edward Island, Ontario, Quebec,
Saskatchewan. Costa Rica. Chile. Region
VIII. Ecuador. Guatemala. Mexico: Guer-
rero, Jalisco, Nuevo Leon, San Luis Potosi,
Tamaulipas. United States: Arizona, Colo-
rado, Florida, Georgia, Illinois, Iowa, Kan-
sas, Maryland, Mississippi, Missouri, Mon-
tana, Nevada, New Mexico, North Carolina,
Oregon, Pennsylvania, Tennessee, Texas,
Utah, Virginia, Washington, Wisconsin,
Wyoming. West Indies.

Hosts. — Coleoptera: Curculionidae (You-
suf and Shafee 1988). Lepidoptera: Cossi-

dae (Pan and Lim 1980), Pyralidae (Schei-
belreiter 1976), Noctuidae (Scheibelreiter
1980).

40. Tutnidifemur Girault

(Figs 58, 133, 134, 160)

Tumidifemur Girault 1911. Type species: Tumidi-
femur pulchrum Girault, by original designa-
tion.

Diagnosis. — Female. Antenna with 2 anelli
and 4 somewhat asymmetrical club seg-
ments; club subcorneal, segments subequal
in length, widest at CI and gradually
narrowing to apex, surface of CI and C2
finely, longitudinally ridged. Fore wing
moderately wide, c. twice as long as wide;
venation with elongate MV, SV angling
only slightly away from wing margin,
directed toward wing apex, PM narrower
than MV with two strong setae, MV and
PM confluent or very narrowly separated,
with several short stiff setae immediately
anterior to PM in costal cell and/or on MV
itself; disk densely setose beyond venation
but often with a large glabrous area near
center, generally without distinct linear
setal tracks although glabrous area may
result in a linear track at its basal border
(resembling an RSI) (cf. Figs 133, 134), also
setal tracks may be distinguishable along
extreme anterior and posterior section of
disk. Hind wing relatively wide, cultri-
form, with length of posterior fringe only
slightly greater than maximum wing
width. Mesotibia with apical spur elongate,
fringed, as long or longer than first
tarsomere.

Male. GC reduced, small, narrowing at
base, without VS, PAR, ventral setae or
aedeagal AAP; GL less than half HTL.

Distribution. — New World tropics and
China.

Diversity. — Two species have been de-
scribed. In the New World, Tumidifemur
has heretofore been known only from the
type series of T. pulchrum from Trinidad
(examined, NMNH). The second named
species occurs in China (see below). A few

Volume 15, Number 1, 2006

113

undescribed species are known from Cen-
tral and South America.

Discussion. — The genus is best separated
by its four-segmented club and unique fore
wing features (Figs 133, 134, 160). The latter
include the stigmal vein slanted only
slightly away from the marginal vein, the
densely setose costal cell and /or marginal
vein, and the densely setose disk which
lacks linear setal tracks in the area apical to
the venation. Tumidifemur is somewhat
anomalous and a close relationship to other
genera is not obvious. The four-segmented,
subcorneal club, tumid femora and highly
reduced male genitalia suggest a tie to
Uscana but the latter lacks the fore wing
traits defining Tumidifemur.

The type species of Tumidifemur has
a patch of dense setation immediately
behind the marginal vein, but this does
not characterize the undescribed species.
Girault (1911b) and Doutt and Viggiani
(1968) characterized the club of Tumidife-
mur as three segmented. Examination of
the type and all additional material
shows it to be four segmented. The
placement of T. ramispinum Lin from China
in this genus is questionable. I have not
examined this species but illustrations in
Lin (1994) indicate a five-segmented club
and genitalia with aedeagal apodemes,
features not characterizing New World
Tumidifemur.

New world records. — Colombia. Costa
Rica. Ecuador. Venezuela. West Indies.

Hosts. — Hemiptera: Membracidae (Gir-
ault 1911b).

41. Ufens Girault

(Figs 59, 60, 135, 238)

Ufens Girault 1911. Type species: Trichogramma
nigrum Ashmead, by original designation.

Because Ufens currently is being revised
by A. K. Owen (UCRC) generic treatment is
abbreviated. Omitted are the complete list of
generic synonyms and specific distribution
records for the genus. None of the omitted
synonyms relate to New World taxa.

Diagnosis. — Female. Antenna with 2 an-
elli, 2 funicular and 3 club segments; Fl
relatively large, subequal in width to F2, all
postanellar segments with PLS and FS.
Fore wing relatively wide (c. 1.7X as long
as wide) with short fringe setae (length c.
0.1 X FWW); venation with PM, MV and SV
subequal in length, radial process present;
disk moderately densely setose with linear
setal tracks including RSI distinct.

Male. Club with a small apical fourth
segment, C4 with FS but not PLS, postanel-
lar segments with conspicuous whorls of FS.
Genitalia extremely variable but ADG never
distinct from GC; aedeagal AAP absent in
New World species; VS, PAR and IVP
usually present and highly modified.

Distribution. — Worldwide except not yet
recorded from South America. In the New
World, Ufens occurs from Canada to
Central America, including the West In-
dies.

Diversity. — There are 30 named species
of Ufens. Some of these are incorrectly
placed and several additions are being
described (Owen, in prep.). Approximately
forty-five species of Ufens now occur in
collections. Three currently are recognized
in the New World (Al-Wahaibi et al. 2005).
These include U. niger (Ashmead), LI.
ceratus Owen, and U. principalis Owen.
Four additional species await description
(Owen, pers. comm.).

Discussion. — Ufens is most easily con-
fused with Mirufens, Zagella and Burksiella.
Wing structure is comparable, and, in
females of all genera, antennal structure is
similar as well (two-segmented funicle,
three-segmented club). In addition, males
of Mirufens have a small fourth club
segment as do Ufens males. Mirufens is
easily separated by its two-segmented
maxillary palp, the transversely ridged
pedicel and the row of spines on the fore
tibia. Zagella and Burksiella are separated
by funicle structure. In Ufens both IT and
F2 are similar in width, and each bears
PLS. In Zagella and Burksiella, Fl is consid-
erably smaller, narrower, and closely ap-

114

Journal of Hymenoptera Research

pressed to F2; it never bears PLS. Besides
Mi ru fe us, the only other genus with a C4 in
males is Ceratogramma. The two-segmented
palp, three anelli and absence of the setal
track RSI in Ceratogramma separate it from
Ufens.

Hosts. — Hemiptera: Cicadellidae (Al-
Wahaibi et al. 2005). Orthoptera: Tettigo-
niidae (Timberlake 1927).

42. Uscana Girault

(Figs 61, 136)

Uscana Girault 1911. Type species: Uscana
semifumipennis Girault, by original designa-
tion.

Bruchoctonus Grese 1923. Type species: Bruchoc-
tonus senex Grese, presumably by monotypy
(see Fursov 1988).

Diagnosis. — Color of mesosoma and me-
tasoma relatively uniform, brownish. Fe-
male. Antenna usually with 2 anellar and 4
club segments; CI often slightly separated
from C2, as long or longer than C2 and
with at least 1 PLS. Midlobe of mesoscu-
tum and scutellum each with 2 pair of setae,
both pairs on each sclerite subequal in
length. Fore wing variable in shape, 1.8-
2.0 X as long as wide in New World species,
fringe setae relatively short, < 0.2 X FWW;
venation with SV distinct, directed toward
posterior margin of wing; disk moderately
densely setose, linear setal tracks (including
RSI) present but not always clearly distin-
guished from adjacent setae.

Male. Antenna with or without PLS on
CI. GC highly reduced, short, without VS,
PAR or ventral setae; aedeagal AAP absent.

Distribution. — Worldwide.

Diversity. — Uscana contains 26 species.
Only three are known from the New
World. Included are Li. semifumipennis
from the United States, and 11. espinae
Pintureau and Gerding and U. chilensis
Pintureau and Gerding, from Chile. Cer-
tain species (U. semifumipennis in particu-
lar) have been broadly introduced for
biological control of bruchid beetles (Fur-
1995b). Although widespread, Uscana

is not commonly collected by standard
bulk sampling methods.

Discussion. — Uscana is not a very distinc-
tive genus and superficially resembles
several others. However, its four-segment-
ed club and lack of a funcle will separate it
from most. In the New World it is only
likely confused with Pseuduscana, Tumidi-
femur and Uscanopsis which have a similar
antennal formula. Pseuduscana is separated
by its considerably shorter CI which lacks
a PLS; the two usually can also be
separated by color (see Pseuduscana). Tu-
midifemur is distinguished by its densely
setose fore wing which lacks setal tracks on
most of the discal surface, as well the
unique characters associated with its vena-
tion. The orientation of the stigmal vein
should also separate Tumidifemur. In Tumi-
difemur it is directed toward the apical
wing margin; in Uscana it is directed
toward the posterior wing margin. The
poorly known Uscanopsis, apparently with
similar antennal structure, is separated by
its exceptionally long hind tibial spur.

Yousuf and Shafee (1988) treated Zaga as
a junior synonym of Uscana. This synony-
my is not accepted (see Zaga below).

New World records. — Brazil: Santa Catar-
ina. Argentina: La Rioja, Misiones. Chile:
(see Pintureau et al. 1999). Guatemala.
Mexico: Baja California Sur, Oaxaca, San
Luis Potosi, Sinaloa, Tamaulipas. United
States: Arizona, California, Georgia, Texas.

Hosts. — Coleoptera: Bruchidae (Fursov
1995b, Pintureau et al. 1999).

43. Uscanella Girault

Uscanella Girault 1911. Type species: Uscanella
bicolor Girault, by original designation.

Diagnosis. — Female: Antenna with 2 anel-
lar and 3 club segments; CI short, discoid,
appressed to C2. Fore wing distinctive:
moderately wide with fringe setae elon-
gate, c. half as long as FWW, apical margin
of wing arcuate; venation with an elongate,
parallel sided SV (i.e. without an expanded
stigma) which is directed toward the apical

Volume 15, Number 1, 2006

115

margin of wing, a short postmarginal vein
present; disk sparsely setose overall with
few linear setal tracks, setation moderately
dense anterior to r-m but subglabrous
between r-m and Cu2, RSI absent; discal
setae extremely short, their sockets more
apparent than the setae themselves. Fem-
ora tumid.

Male. Unknown.

Distribution. — Known only from Trini-
dad and Tobago.

Discussion. — Uscanella is known only
from the original types (examined, NMNH).
The three-segmented club, short and trans-
verse CI, distinctive stigmal vein and the
very short, relatively sparse setation on the
fore wing separate it from other genera. The
orientation and shape of the stigmal vein are
somewhat similar in Viggianiella, although
its length is considerably shorter in Usca-
nella; also the two are dissimilar for most
other features. Fore wing structure also
resembles that in Pteranomalogramma but
the apical margin is arcuate in Uscanella,
not pointed. Also the structure of the stigmal
vein differs considerably. Because males are
unknown, Uscanella is tentatively placed in
the Chaetostrichini based on antennal struc-
ture. The genus is illustrated by Doutt and
Viggiani (1968).

Records. — West Indies.

Hosts. — Hemiptera: Membracidae (Gir-
ault 1911b).

44. Uscanoidea Girault

(Figs 78, 147, 182, 250)

Uscanoidea Girault 1911. Type species: Uscanoi-
dea nigriventris Girault, by original designa-
tion.

Gnorimogramma De Santis 1972. Type species:
Gnorimogramma aliciae De Santis, by original
designation. New synonymy.

Diagnosis. — Female. Antenna with 2 anelli
and 5 asymmetrical club segments, club
widest at C2, C2 often loosely associated
with C3, C3 without APB sensilla. Fore
wing 1.6-1 .9 X as long as wide, fringe setae
0.1 X FWW; venation extending 0.4-0.5

FWT, SV constricted at base and extending
apically beyond apex of MV, MV 1.4-1.6X
(uncommonly 1.2 X) as long as PM and
subequal in width, SV+MV/PM >1.4 (usu-
ally 1.7-2.0); disk moderately densely
setose, with distinct linear setal tracks,
RSI elongate with more than 5 setae,
broadly curved from apex of stigma to-
ward base of wing where it converges on
the Cu tracks. Propodeal disk length
subequal to that of metanotum (Fig. 182).
Fore tibia rarely spinose dorsally but if so
then entire surface spinose and not with
a single spine at middle more prominent
than others. Ovipositor variable in length
but never extending appreciably beyond
apex of metasoma.

Male. — Antennal club segments usually
less asymmetrical than in female. GC
gradually narrowed to apex, rounded at
base, usually without PAR, ventral setae
and aedeagal AAP; GL considerably less
than HTL (see Discussion).

Distribution. — Known from the Oriental,
Nearctic and Neotropical regions. In the
New World, distributed from the United
States to Argentina. Uncommonly collected
north of Mexico.

Diversity. — Twelve species are assigned
to Uscanoidea. Included are five Chinese
species [three described as Gnorimogramma:
U. acuminate (Lin), U. aduncata (Lin) and
Li. oviclavata (Lin), new combinations], and
seven from the New World. Six of the latter
were treated by Viggiani (1992) and De
Santis (1997). The New World species
include U. aliciae (De Santis), new combi-
nation (from Gnorimogramma); U. hastata
De Santis; U. iperterebrata Viggiani; U.
marilandica (Girault); U. nigriventris Girault;
U. parviclavata De Santis; U. silvestrii
Viggiani. U. marilandica is North American;
the remainder occur in Central and South
America. As with Burksiclla and Lathromer-
oidea there are numerous undescribed
species of Uscanoidea, particularly in the
neotropics. Placement of U. marilandica
here is based on Viggiani (1986, 1992) and
notes taken from the type (NMNH) over 15

116

Journal of Hymenoptera Research

years ago. Attempts were made to confirm
generic placement for this study, however
the type has been on loan for over a decade
and attempts at recall by NMNH were not
successful.

Discussion. — Uscanoidea is defined here
as having an antennal structure similar to
Zaga but fore wings as in Burksiella. Also,
unlike Zaga, ventral spines on the genital
capsule are almost always absent in Usca-
noidea. Male and female syntypes of U.
nigriventris (examined, NMNH) are in very
poor condition but diagnostic features of
the genus are visible.

Gnorimogramma, synonymized here with
Uscanoidea, was compared to Chaetostricha
by De Santis (1972) and separated by the
narrower club and shorter ovipositor. It
also differs from Chaetostricha in lacking
both a distinct funicle and a single promi-
nent spine on the fore tibia, by its wider
fore wing with an arcuate RSI, and its
considerably shorter genital capsule. These
features place it in Uscanoidea as herein
defined. The antennal club and ovipositor
characteristics cited by De Santis to estab-
lish Gnorimogramma vary widely within
most genera of the Chaetostricha group.
Paratypes (UCRC) of Gnorimogramma ali-
ciae, the type species, were examined.

The separation of Uscanoidea species
from Group B of Lathromeroidea is not
always straightforward. In a few species
the wings are characteristic of the latter
(narrower, longer fringe) but, as in Usca-
noidea, the propodeum is not much, if at all,
longer than the metanotum. These cannot
be placed unambiguously based on current
generic limits.

Uscanoidea also is similar to Centrobiopsis.
It differs primarily by its broader fore wing,
and shorter ovipositor and male genitalia.
Unlike Centrobiopsis, in Uscanoidea the ovi-
positor does not extend beyond the meta-
soma and the genital length is considerably
less than that of the hind tibia.

New World records.— Argentina: Chaco,
Formosa, La Rioja, Misiones, Salta, San-

tiago del Estero. Belize. Brazil: Rio
de Janeiro, Mato Grosso do Sul, Santa
Catarina, Sao Paulo. Colombia. Costa
Rica. Ecuador. Guatamala. Mexico: Chia-
pas, Colima, Nuevo Leon, Sinaloa, Tamau-
lipas, Veracruz, Yucatan. Nicaragua. Pan-
ama. Peru. United States: Florida, Mary-
land, Oklahoma, South Carolina. West
Indies.

Hosts. — Hemiptera: Cercopidae (De San-
tis 1972), Cicadellidae (unpubl. record),
Membracidae (De Santis 1997).

45. Uscanopsis Girault

(Fig. 137)

Uscanopsis Girault 1916. Type species: Uscanop-
sis carlylei Girault, by original designation.

Diagnosis. — Female. Antenna apparently
with a 4-segmented club (but see Discus-
sion), club subcorneal, broadest at base.
Propodeal disk triangular, distinctly longer
than metanotum. Fore wing c. 2.2 X as long
as wide; venation confluent, PM c. 3x
wider than MV, MV gradually curving
away from wing margin to form SV, SV
basal to stigma subequal in width to MV;
disk moderately densely setose, all setae
scattered and not arranged in linear tracks,
RSI absent. Femora tumid; tarsi short,
robust, metatarsus only about half HTL;
hind tibial spur unique, extremely elongate
and stout, c. 0.9 X length of metatarsus,
dorsal surface of spur (surface facing
tarsus) microserrate.

Male. GC with aedeagal AAP present
(presence of other genital structures ques-
tionable).

Distribution. — Known only from the type
locality, Port of Spain, Trinidad and To-
bago.

Diversity. — Monotypic.

Discussion. — Uscanopsis is known only
from the type series of U. carlylei (exam-
ined, NMNH). All specimens, on a single
slide, are in very poor condition and
several characters cannot be adequately
viewed. Girault (1916b) considered the
antennal club as two segmented; Doutt

Volume 15, Number 1, 2006

117

and Viggiani considered it four segmented.
I also detect four club segments, but there
remains the possibility of a very small first
segment as occurs in Uscanoidea and other
genera which, if present, is impossible to
see in the types. Fortunately, the identifi-
cation of Uscanopsis is assured by the very
long and stout hind tibial spur which has
no counterpart in the family. Also fore
wing structure appears to be characteristic
(Fig. 137). The veins are confluent, the
premarginal is considerably wider than the
marginal vein, and the latter gradually
curves apically to form the stigmal vein.
The considerably setose wing disk without
linear setal tracks also helps characterize
the genus.

Uscanopsis is not obviously related to any
other group. A wide premarginal relative
to the marginal vein also occurs in Adryas,
but as indicated by Pinto and Owen (2004),
the shape of the premarginal differs in the
two and this is accompanied by several
other differences as well. Better preserved
specimens are required before comparing
Uscanopsis with other elements in the
family.

The drawings of antenna, fore wing,
maxilla, and hind leg of Uscanopsis in
Doutt and Viggiani (1968, Fig. 46) are
useful for identification, however my ex-
amination of the types suggest certain
corrections. In their drawing of the fore
wing the marginal vein is broader than in
the types. Their Fig. 46D shows the apex of
the marginal as wider than the base of the
stigmal vein, whereas it is equally wide.
Also, I find the hind tibial spur of the male
syntype to be longer than that illustrated
by Doutt and Viggiani (Fig. 46C). The
illustration of the antenna (Fig. 46A) shows
the apex of the club as deeply bifid.
Although indeed the case in at least one
of the syntypes, this may simply be due to
shrinkage of cuticle between the two apical
PLS during slide preparation.

Records. — West Indies.

Hosts. — Hemiptera: Membracidae (Gir-
ault 1916b).

46. Xiphogramma Nowicki

(Figs 62, 138, 195, 239, 240)

Xiphogramma Nowicki 1940. Type species: Xi-
phogramma holorhoptra Nowicki, by original
designation. Pinto 1990b (generic review, key
to spp.).

Diagnosis. — Female. Antenna with 2 anelli,
2 funicular and 1 club segment, funicular
segments similar in length and width; PLS
present on funicle, numerous short and fine
APB sensilla on club. Fore wing with base of
MV and apex of PM poorly sclerotized, light
in color, SV short, lacking a basal constric-
tion, MV without numerous heavy setae
dorsally; disk densely setose but dense
setation not extending basally behind vena-
tion, linear setal tracks poorly indicated at
least apically, RSI absent. Ovipositor elon-
gate, extending length of metasoma and
with apical 0.3-0.4 reaching beyond meta-
somal apex.

Distribution. — Europe, Asia, Africa,
North America.

Diversity. — Four species of Xiphogramma
are described. Included are X. nnneckei
Doutt (Africa), X. holorhoptra (Europe), X.
indicum Hayat (Asia) and X. fuscum Pinto
(North America).

Discussion. — A discussion of generic lim-
its and separation of Xiphogramma from the
closely related Chaetogramma and Brachy-
grammatella is provided in the treatment of
Chaetogramma. Xiphogramma fuscum, the
only New World representative, is sepa-
rated from Chaetogramma by ovipositor
length. It is further separated from New
World Chaetogramma by the completely
divided funicular segments, the shape of
the genital capsule (attenuate at base in X.
fuscum, truncate basally in Chaetogramma),
and the absence of aedeagal apodemes.
These features do not separate the genus
from Old World Chaetogramma however.

New World records. — Canada: Alberta.
Mexico: Baja California Sur, Sinaloa, So-
nora. United States: Arizona, California,
New Mexico, Oregon, Texas, Utah.

Hosts. — Unknown.

118

Journal of Hymenoptera Research

47. Zaga Girault, renewed status

(Figs 81-83, 149, 162, 177, 252, 253)

Zaga Girault 1911. Type species: Zaga latipennis
Girault, by original designation. Yousuf and
Shafee 1988 (as junior synonym of Uscana
Girault).

Lathrogramma De Santis 1952. Type species:
Lathrogramma deltae De Santis, by original
designation. New synonymy.

Diagjiosis. — Female. Antenna with 2 anelli
and 5 club segments, club segments asym-
metrical, CI very short and closely ap-
pressed to C2, C2 lined longitudinally and
with several PLS, C3 without APB sensilla.
Fore wing broad, oblate, 1. 5-2.0 X as long
as broad, fringe setae very short, < 0.1 X
FWW (usually < 0.05); venation short, not
exceeding 0.5 FWL, MV short, wide, c.
0.4 X as wide as long and subequal in
length to and considerably wider than PM,
SV+MV/PM < 1.5 (usually < 1.3); SV not
constricted basally, not extending beyond
apex of MV (a line drawn through SV to
wing margin describes a right angle with
MV); disk moderately densely setose, with
distinct linear setal tracks, RSI present and
usually arranged in a straight line sub-
perpendicular to Cu tracks, less commonly
curving posteriorly toward base of wing.
Fore tibiae usually with spine(s) on dorsal
surface, most prominent spine near middle
as in Qiaetostricha (Fig. 177).

Male. GC similar to that in Burksiella and
Uscanoidea but almost always with ventral
setae.

Distribution. — Known only from Hawaii
and the New World where it has been taken
from Canada S to Ecuador; also present in
Argentina.

Diversity. — Only two named species are
assigned to Zaga, Z. latipennis, the type
species from Virginia, and Z. deltae (De
Santis), new combination (from Lathro-
gramma), from Argentina. Several unde-
scribed species occur in North and South
America.

Discussion. — Zaga is separated from sim-
sra by its antennal formula and fore

wing structure. It is the only genus lacking
a funicle and with a five-segmented club in
which the stigmal vein is not constricted
basally and is oriented perpendicular to
the marginal vein (Figs 149, 162). Only
Zagella, which has a distinct funicle, has
similar venation and it is likely that the two
are closely related. In Uscanoidea, also with
a five-segmented club, the marginal vein is
more slender and the stigmal vein is
extended apically describing an oblique
angle with the marginal vein (cf. Figs 161,
162).

Lathrogramma, herein treated as a syno-
nym of Zaga, was considered close to the
Old World trichogrammatine Ophioneuris
by De Santis (1952) based on superficial
similarity in fore wing structure. The
assumption that the club was only four
segmented in Zaga (Girault 1911a, Doutt
and Viggiani 1968) led to its being com-
pared to Uscana rather than Lathrogramma.
This assumption prompted Yousuf and
Shafee (1988) to synonymize Zaga with
Uscana. I examined types of Zaga latipennis
(NMNH) and Lathrogramma deltae (para-
type, UCRC) and have verified that the
club is similarly five segmented in both. As
in other genera of the Qiaetostricha group,
CI is very small and sometimes difficult to
discern.

Z. latipennis and Z. deltae differ consid-
erably in size and ovipositor length. The
former is among the largest of trichogram-
matids with a long ovipositor spanning the
entire length of the metasoma. Z. deltae, on
the other hand, is much smaller with
a considerably shorter ovipositor. The
structure of the wings, antennae and male
genitalia are similar in both, and differ-
ences in ovipositor length are bridged by
undescribed species. Most of the unde-
scribed species resemble Z. deltae more
closely.

New World records. — Argentina: Buenos
Aires. Bermuda. Canada: British Colum-
bia, Ontario. Brazil: (R. B. Querino, pers.
comm). Costa Rica. Ecuador. Guatemala.
Honduras. Mexico: Baja California Norte,

Volume 15, Number 1, 2006

119

Baja California Sur, Chiapas, Colima, Ja-
lisco, Morelos, Nuevo Leon, Oaxaca, Quer-
etero, Quintana Roo, San Luis Potosi,
Sinaloa, Veracruz, Yucatan. United States:
Arizona, California, Florida, Georgia, Illi-
nois, Kansas, Maryland, Missouri, Nevada,
New Hampshire, New Mexico, North
Carolina, Oklahoma, Oregon, South Car-
olina, Tennessee, Texas, Utah, Virginia,
West Virginia, Wyoming. West Indies.
Hosts. — Unknown.

48. Zagella Girault

(Figs 5, 84, 85, 150, 254-257, 260)

Zagella Girault 1918. Type species: Paracentrobia
flavipes Girault, by original designation. Tria-
pitsyn 2003 (lectotype designation for type
species).

Diagnosis. — Female. Body usually rela-
tively robust. Antenna with 2 anelli, 2
funicular and 3 club segments; Fl short,
anelliform, closely appressed to F2, F2
slightly narrower than CI, with 1 to several
PLS; club segments relatively symmetrical
(i.e., segment length similar on all sur-
faces), CI with APB sensilla. Fore wing
broad, oblate, <2X as long as wide, fringe
setae short, <0.1X FWW; venation short,
not attaining 0.5 wing length, MV short,
broad, < 1.5X length of PM and consider-
ably wider (commonly c. 1.5X width of
PM), PM and base of MV lightly sclero-
tized, SV not constricted at base and not
extending beyond apex of MV (a line drawn
through stigma to wing margin describes
a right angle with MV); SV+MV/PMV < 1.5
(usually c. 1.3), radial process absent; disk
moderately densely setose, with distinct
linear setal tracks, RSI present and arranged
in a straight line, directed toward posterior
margin of wing, subperpendicular to Cu
tracks, not curving posteriorly toward base
of wing, commonly composed of relatively
few setae (usually 5 or fewer). Fore tibia
without spines on dorsal surface. Oviposi-
tor not extending beyond metasomal apex.

Male. GC short, robust, its length less
than HTL, with ventral setae; aedeagal

AAP absent. Genital structure variable: GC
constricted at middle, without VS or PAR
and with basal margin transverse in North
American species (Figs 254, 255); GC with
a pair of modified VS (or PAR?) and basal
margin transverse or emarginate in South
American species (Figs 256, 257).

Distribution. — Confined to the New
World although not common in tropics.

Diversity. — Five species are placed in
Zagella. Besides the type species, Z. flavipes
(Girault), described from eastern United
States, the only named species in the genus
are from temperate South America. Includ-
ed are Z. delicata De Santis, Z. mimica De
Santis, Z. nanula De Santis, and Z. zebrata De
Santis. Several undescribed species are
known from North America especially in
the Southwest. I have examined type
material of all species except Z. nanula and
Z. mimica; the latter two are unknown to me
and placement in Zagella needs confirma-
tion.

Discussion. — The definition of Zagella
adopted here varies from previous treat-
ments. Doutt and Viggiani (1968) consid-
ered Burksiella a junior synonym of Zagella.
Burksiella is herein resurrected and certain
species previously treated as Zagella are
transferred there (see above). Zagella shares
its distinctive fore wing venation with
Zaga. In both the stigmal vein lacks a basal
constriction and is perpendicular to the
marginal vein. They also share a relatively
short and broad marginal vein relative to
the premarginal. The two are separated by
antennal formula (two-segmented funicle,
three-segmented club in Zagella; funicle
absent, five-segmented club in Zaga). Za-
gella is likely confused with Burksiella.
Characters for separation are reviewed in
the treatment of the latter genus.

Zagella appears to have a largely bipolar
distribution in the New World, occurring
in North America and temperate South
America. The only material I have seen
from intermediate areas is a single female
from Venezuela. The South American
species may represent a distinct lineage.

120

Journal of Hymenoptera Research

This difference is indicated by male geni-
talia. In North American species the genital
capsule is simple without obvious para-
meres or volsellae (Fig. 255) whereas the
South American species examined bear
a pair of laterally curved sickle-shaped
structures which either represent modified
parameres or volsellae (Figs 256, 257).

The species described as Bracln/ia radialis
by De Santis (1997) may be a Zagella
judging from illustrations in the original
description. It clearly is not a Brachyia.
Unfortunately the type(s) of this species
can not be located (M. Loiacono, MLPA,
pers. comm.). Although presumably neo-
tropical, De Santis does not indicate a type
locality for this species.

The types of Z. flavipes were studied by
Triapitsyn (2003), who designated a lecto-
type. The included drawing of the male
genitalia, based on one of the paralecto-
types, is misleading. It shows the genital
capsule as evenly tapering from base to
apex and with a rounded basal margin.
These features suggest Burksiella, the genus
most easily confused with Zagella. I have
examined the paralectotype drawn by
Triapitsyn (NMNH), and although the
genitalia are difficult to see well, they are
typical of other North American Zagella,
i.e., the basal margin is transverse and the
capsule is constricted before the apex (as in
Figs 254, 255).

Records. — Argentina: Buenos Aires, Cata-
marca, Chaco, Formosa, La Rioja, Mendosa,
Misiones, Salta, Tucuman. Brazil: Minas
Gerais, Sao Paulo. Mexico: Baja California
Sur, Jalisco, Michoacan, Sinaloa, Sonora,
Tamaulipas, Yucatan, Zacatecas. Uruguay.
Venezuela. United States: Arizona, Califor-
nia, Florida, Georgia, Maryland, New Mex-
ico, Oregon, Texas, Utah, Washington.

Hosts. — Hemiptera: Cicadellidae (Lo-
garzo et al. 2004).

Oligositini Walker

Diagnosis. — Eyes black. Antenna with no
more than 4 postanellar segments; funicle

usually present, always 1 segmented
(Figs 86-93). One pair of setae on both
midlobe of mesoscutum and on scutellum
(Figs 186, 187). Type 2B genitalia (Figs 258,
259): GC with ADA extremely reduced and
largely directed anteriorly, VS, PAR, ven-
tral setae and aedeagal AAP always absent.
Discussion. — Because the tribe was recent-
ly reviewed by Pinto and Viggiani (2004) an
abbreviated treatment is presented here.
They divided the group into two subtribes,
Oligositina and Eteroligositina, presented
a preliminary hypothesis of generic relation-
ships and a key to the World genera.

Oligositini: Oligositina Walker

Diagnosis. — Propodeal disk usually dis-
tinctly longer than metanotum at midline
(Fig. 186). Mesopleuron with pleural suture
(Fig. 192); transepisternal sulci absent. Meta-
soma with terga uniformly sclerotized their
entire length. Male genitalia highly reduced
to a simple tube-like structure (Fig. 258).

Discussion. — As noted by Pinto and
Viggiani (2005), support for the monophyly
of this subtribe is relatively weak.

49. Epoligosita Girault

(Figs 86, 151)

Paroligosita Girault and Dodd, in Girault 1915
(as subgenus of Oligosita). Type species:
Paroligosita biclavata Girault and Dodd, by
original designation.

Epoligosita Girault 1916 (as genus; ;;. n. for
Paroligosita Girault and Dodd, nee Kurdjumov
1911).

Epoligositina Livingstone and Yacoob 1983 (as
subgenus). Type species: Epoligosita (Epoligo-
sita) duliniae Livingstone and Yacoob, by
original designation. Lin 1990 (as genus).
Pinto and Viggiani 2004 (renewed status as
subgenus).

Diagnosis. — Antennal segmentation vari-
able, 1 funicular segment present or absent,
club 1 or 2 (rarely 3) segmented. Tarsi
elongate, fore and middle tarsi distinctly
longer than tibia. Propodeal disk sub-
tended by a small subtriangular lobe. Fore

Volume 15, Number 1, 2006

121

wing c. 3-4 X as long as wide, widest near
level of stigma; venation with PM abruptly
wider than MV at junction; disk glabrous,
with one or two setae at most.

Distribution. — The nominate subgenus is
widespread. Epoligositina has not been
recorded from the New World.

Diversity. — Epoligosita contains 22 species.
Only one, E. mexicana Viggiani, occurs in the
New World. All collections examined from
North and South America either represent
this species or are very similar forms.

Discussion. — Epoligosita is not easily con-
fused with any other genus. The fore wing
alone serves for identification. It is the only
oligositine with a moderately narrow fore
wing (3-4 X as long as wide) which is
essentially glabrous. Some species of Oli-
gosita have a sparsely setose fore wing, but
the three segmented club easily separates
them from Epoligosita, at least in the New
World. Although antennal formula is quite
variable in Epoligosita when considering
the world fauna (Pinto and Viggiani 2004),
all known representatives from North and
South America have a single funicle seg-
ment and a single club segment. At most
the club shows a partial division into two
segments.

New World records. — Argentina: Chaco,
Formosa, Salta. Costa Rica. Ecuador. Gua-
temala. Mexico: Baja California Sur, Coa-
huila, Durango, Guerrero, Sinaloa. United
States: Arizona, California, Texas. West
Indies.

Hosts. — Hemiptera: Cicadellidae (Vig-
giani 1985, Pinto and Viggiani 1987),
Tingidae (Livingstone and Yacoob 1983).

50. Megaphragma Timberlake

(Figs 87, 152)

Megaphragma Timberlake 1923. Type species:
Megaphragma mymaripenne Timberlake, by
original designation.

Sethosiella Kryger 1932. Type species: Sethosiella
priesneri Kryger, by original designation.

Paramegapliragma Lin 1992. Type species: Para-
megapliragma stenopterum Lin, by original
designation. Delvare 1993 (synonymy).

Diagnosis. — Extremely small, body leng-
th <0.3 mm.

Antenna with or without a single funicle
segment, club 2 or 3 segmented, never with
more than 3 postanellar segments (i.e. if
club 3 segmented then funicle absent).
Maxillary palp small but present. Fore
wing extremely narrow, strap-shaped, c.
7x as long as wide; disk glabrous or with 1
or 2 rows of few setae. Metasomal tergum
VII without spiracles.

Distribution. — Widespread.

Diversity. — Fifteen species are assigned
to Megaphragma. Only three, M. mymarip-
enne (widespread), M. striatum Viggiani
(Mexico) and M. caribea Delvare (Guade-
loupe), occur in the New World.

Discussion. — The genus is distinguished
by its very small body size, extremely
narrow, strap-like fore wings, antennae
with three or fewer postanellar segments,
and absence of metasomal spiracles. Its
closest relatives appear to be Prestwiclua
and Sinepalpigramma (Pinto and Viggiani
2004).

New World records. — Argentina: Buenos
Aires, Formosa, Misiones, Salta. Costa
Rica. Mexico: Chiapas. United States:
California. West Indies.

Hosts. — Thysanoptera: several taxa (see
Noyes 2001)/

51. Oligosita Walker

(Figs 88, 89, 153, 168, 172, 186, 192, 258)

Oligosita Walker 1851. Type species: Oligosita

collina Walker, by monotypy.
Westwoodella Ashmead 1904. Type species:

Oligosita subfasciata Westwood, by original

designation.
Paroligosita Kurdjumov 1911. Type species:

Paroligosita bella Kurdjumov, by original

designation.

Diagnosis. — Antenna with 1 funicular
and 3 club segments, funicle distinctly
separated from club, PLS present on club.
Maxillary palp present. Fore wing disk
sparsely to moderately densely setose, no
more than 4X as long as wide. Propodeal

122

Journal of Hymenoptera Research

disk not subtended by a small subtriangu-
lar lobe.

Distribution. — Worldwide.

Diversity. — Forty-six of the 144 species
previously assigned to Oligosita were re-
cently transferred to Pseudoligosita by Pinto
and Viggiani (2004). Of the 98 species
remaining in Oligosita only seven are from
the New World. These include O. americana
Girault, O. clarimaculosa (Girault), O. san-
guined (Girault) and O. subfasciatipennis
(Girault) from North America; O. desantisi
Viggiani from South America; and O.
giraulti Crawford and O. magnified Dozier
from the West Indies. A large number of
undescribed species undoubtedly occur.

Discussion. — All of the described New
World Oligosita belong to the collina Group
as defined by Viggiani (1976b). This as-
semblage is recognized and separated from
congeners and all related genera by the
characteristic clavate apical placoid sensil-
lum on the antennal club of females
(Figs 88, 89). This PLS resembles the
terminal process in certain chaetostrichine
genera. In the latter, however, it is the apex
of the club itself which is prolonged, not
the PLS.

New World records. — Argentina: Buenas
Aires, Formosa, La Rioja, Salta, Tucuman.
Belize. Bolivia. Brazil: Amazonas, Para,
Pernambuco. Canada: Alberta, British Co-
lumbia, Manitoba, Ontario, Prince Edward
Island. Chile: Region Metropolitana. Co-
lombia. Costa Rica. Ecuador. Guatemala.
Mexico: Baja California Sur, Campeche,
Chiapas, Colima, Distrito Federal, Guer-
rero, Jalisco, Morelos, Nuevo Leon, Quin-
tana Roo, Sinaloa, Tamaulipas, Veracruz,
Yucatan. Panama. United States: Arizona,
California, Colorado, Florida, Georgia,
Iowa, Illinois, Kansas, Louisiana, Mary-
land, Michigan, Mississippi, Missouri,
Montana, Nebraska, Nevada, New Mexico,
North Carolina, Oklahoma, Oregon, Penn-
sylvania, South Carolina, South Dakota,
Tennessee, Texas, Utah, Virginia, Wash-
ington, Wisconsin, Wyoming. Venezuela.
West Indies.

Hosts. — Hemiptera: (primarily Cicadelli-
dae, see Noyes 2001).

52. Prestwichia Lubbock

(Figs 90, 154)

Prestwichia Lubbock 1864. Type species: Pres-
twichia aquatica Lubbock, by monotypy.

Austromicron Tillyard 1926. Type species: Aus-
tromicron zi/gopterorwn Tillyard, by original
designation. Doutt and Viggiani 1968 (syn-
onymy).

Diagnosis. — Antenna with 1 funicular
and 3 club segments, funicular segment
closely associated with club (club often
appearing 4 segmented); antenna with PLS
on surface of segments. Maxillary palp
present. Fore wing narrow, c. 7x as long as
wide; disk densely setose in apical half,
setae not arranged in linear tracks. Meta-
somal tergum VII without spiracles. Males
commonly wingless.

Distribution. — Australia, Asia, Europe,
Africa and North America.

Diversity. — Five species of Prestzvichia
have been described. Two undetermined
collections from Florida are the only re-
cords from the New World.

Discussion. — The narrow, highly setose
fore wing and antennal structure separate
Prestwichia from other oligositines. It is most
likely confused with Sinepalpigramma of
Central and South America. Antennal struc-
ture is superficially similar in the two genera
but Sinepalpigramma lacks PLS on the anten-
nal surface as well as maxillary palpi; also its
fore wings are considerably wider.

New World records. — United States: Flor-
ida.

Hosts. — Prestzvichia occurs in aquatic ha-
bitats where hosts of a variety of aquatic
insect groups (Coleopera, Hemiptera, Odo-
nata) are parasitized (Fursov 1995c).

53. Sinepalpigramma Viggiani and Pinto

(Figs 91, 155, 171)

Sinepalpigramma Viggiani and Pinto 2003. Type
species: Sinepalpigramma lougiciliatum Vig-
giani and Pinto, by monotypy.

Volume 15, Number 1, 2006

123

Diagnosis. — Antenna with 1 funicular
segment and a 3-segmented club, funicle
closely associated with club; antenna with-
out PLS. Maxillary palp absent. Surface of
mesoscutum and scutellum smooth, with-
out indication of- reticulae. Metasomal
tergum VII without spiracles.

Distribution. — From northern Mexico to
Argentina.

Diversity. — Two species of Sinepalpi-
gramma have been described, the type
species and S. longiterebratum Viggiani
and Pinto. S. longiciliatum, is the more
widespread occurring throughout the
range of the genus. S. longitei'ebratwn is
known only from Ecuador. A possible
third species occurs in Brazil (Viggiani
and Pinto 2003).

Discussion. — The genus is easily recog-
nized by the absence of maxillary palps.
Overall antennal structure is similar to
Prestwichia but unlike that genus the
antennal surface lacks PLS; also the fore
wings are considerably wider in Sinepalpi-
gramma. Males have not been clearly
associated.

New World records. — Argentina: Mis-
iones. Brazil: Minas Gerais. Colombia.
Ecuador. Costa Rica. Mexico: Michoacan,
Nuevo Leon, Tamaulipas. Panama. Vene-
zuela.

Hosts. — Unknown.

Oligositini: Eteroligositina Lin

Diagnosis. — Propodeal disk usually only
slightly longer than metanotum (Fig. 187);
mesopleuron without a pleural suture
(Fig. 197); transepisternal sulci present;
metasoma almost always with at least
anterior 3 terga longitudinally striate pos-
teriorly (Figs 187, 197). Male genitalia more
variable than in Oligositina.

Discussion. — Unlike the Oligositina, the
monophyly of the Eteroligositina is on
firmer ground (Pinto and Viggiani 2004).
Six genera are assigned to this subtribe
(Pinto and Viggiani 2004). Only two,
Doirania and Pseudoligosita, occur in the

New World. The extralimital genera are
Chaetostrichella, Eteroligosita, Hayatia, and
Probrachista.

54. Doirania Waterson
(Figs 92, 156)

Doirania Waterson 1928. Type species: Doirania
leefmansi Waterson, by original designation.
Pinto 2004 (generic review).

Diagnosis. — Antenna with 1 funicular
and 1 club segment, funicle well separated
from club, transverse. Metasoma with
anterior 3 terga longitudinally striate pos-
teriorly. Male genitalia simple, reduced to
a single tube with two short apodemes at
base.

Distribution. — North America north of
Mexico; Palearctic Region, Indonesia and
New Guinea. A record from Ecuador
(Pinto 2004) requires confirmation.

Diversity. — There are three species of
Doirania. The only New World species is
D. elegans Pinto.

Discussion. — Doirania, as currently de-
fined, is easily separated from other oligo-
sitine genera. The one-segmented funicle
and club, the transverse funicular segment
along with the longitudinally striate meta-
somal terga are characteristic. It is closely
related to Pseudoligosita (Pinto and Viggiani
2004), a genus with three club segments
and a usually longer funicular segment. A
few species with three club segments and
identifiable as Pseudoligosita on this basis
may actually belong to Doirania, which
would alter its definition (see Pinto 2004).
These forms are known only from females
and the absence of males precludes clear-
cut placement at the present time. In males
of Doirania the genitalia are extremely
reduced as in the Oligositina (as in
Fig. 258). In Pseudoligosita the base of the
genitalia have distinctive posteriorly di-
rected apodemes (Fig. 259).

Neiv World records. — Occurring in the
eastern United States and southeastern
Canada, with a single record from Arizona.
See Pinto (2004) for specific locales.

124

Journal of Hymenoptera Research

Hosts. — Orthoptera: Tettigoniidae (Ca-
udwell 2000).

55. Pseudoligosita Girault
(Figs 93, 157, 197, 259)

Pseudoligosita Girault 1913. Type species: Pseu-
doligosita amoldi Girault, by original designa-
tion. Pinto and Viggiani 2004 (renewed
status).

Zorontogramma Silvestri 1915. Type species:
Zorontogramma distinction Silvestri, by origi-
nal designation. Doutt and Viggiani 1968 (as
subgenus of Oligosita). Pinto and Viggiani
2004 (synonymy).

Diagnosis. — Distinguished from Doirania,
the only other eteroligositine genus in the
New World, by the 3-segmented club. Fore
wing disk usually with a crescent-shaped
fumation at apex of stigma, apex of
marking directed toward base of wing.
Male genitalia strongly curved ventrally
with posteriorly directed apodemes at base
(Fig. 259).

Distribution. — Worldwide.

Diversity. — Forty-six species were trans-
ferred from Oligosita to Pseudoligosita by
Pinto and Viggiani (2004). Of these, five
occur in the New World. Included are P.
cosmosipennis (Girault), P. marilandia (Gir-
ault) and P. plebia (Perkins) from North
America; and P. fasciata (Viggiani) and P.
longifrangiata (Viggiani) from South Amer-
ica. These species are not easily identifiable
but it is clear that several undescribed
forms remain.

Discussion. — Although formerly consid-
ered members of Oligosita, the species of
Pseudoligosita have a closer relationship to
the Old World Chaetostrichella and Probra-
chista. As in all Eteroligositina the anterior-
most visible metasomal terga are divided
into an anterior uniformly sclerotized
portion and a posterior, longitudinally
striate section. In the New World only
Doirania has a similar feature. In Doirania
the club is one segmented, and the male
genitalia lack the posteriorly directed apo-
demes at the base.

New World records. — Argentina: Chaco,
Formosa, Jujuy, La Rioja, Misiones, Salta,
Tucuman. Bolivia. Brazil: Minas Gerais,
Sao Paulo. Canada: Alberta, Manitoba, New
Brunswick, Nova Scotia, Quebec. Colom-
bia. Costa Rica. Ecuador. Guatemala. Hon-
duras. Mexico: Baja California Norte, Baja
California Sur, Campeche, Chiapas, Guer-
rero, Jalisco, Michoacan, Morelos, Nuevo
Leon, Oaxaca, Quintana Roo, San Luis
Potosi, Sinaloa, Sonora, Tamaulipas, Vera-
cruz, Yucatan. Panama. United States:
Arizona, California, Colorado, Florida,
Georgia, Illinois, Kansas, Maryland, Michi-
gan, Mississippi, Missouri, New Mexico,
North Carolina, Oklahoma, Pennsylvania,
South Carolina, South Dakota, Tennessee,
Texas, Virginia, West Virginia, Wisconsin.
Uruguay. Venezuela. West Indies.

Hosts. — Hemiptera, Orthoptera, and Co-
leoptera (see Pinto and Viggiani 2004).

ACKNOWLEDGMENTS

This work owes a great debt of gratitude to the
collection managers and collectors who provided
specimens for study. This consisted of important
curated material and type specimens as well as
a wealth of uncurated material gleaned from numer-
ous bulk samples from throughout the Western
Hemisphere. The following individuals and their
current or former institutions are acknowledged in
this regard: John Noyes (BMNH), John Huber (CNC),
Gennaro Viggiani (DEZA), Robert Zuparko (CAS,
EMEC), Michael Sharkey (HIC), Colin Favret (INHS),
Arturo Roig-Alsina (MACN), Paul Hanson (MZCR),
Marta Loiacono and the late Luis De Santis (MLPA),
Terry Erwin, Michael Gates and Michael Schauff
(NMNH), James Woolley (TAMU), Steve Heydon
(UCDC), Jeremiah George, Gordon Gordh, John
Heraty, John LaSalle, James Munro, Albert Owen,
Serguei Triapitsyn, and Doug Yanega (UCRC).

Special acknowledgment is owed to Gary Platner
for preparing most of the slide-mounted material
used in this study, for considerable assistance with
the SEM work, for plate preparation and for
providing additional help in all phases of the project.
Rob Velten was influential in developing the
protocol used for slide mounting and also helped in
curating material. Drawings were prepared by Marina
Planoutene.

Financial support for this study was provided by
a grant from the USDA (NRIICGP) (2001-35316-11012;
J. Pinto, PI), an NSF PEET grant (BSR-9978150; J.
Heraty, PI), and my department at U.C. Riverside.

Volume 15, Number 1, 2006

125

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Kansas Entomological Society 67: 301-310.
Velten, R. K. and J. D. Pinto. 1990. Soikiella Nowicki
(Hymenoptera: Trichogrammatidae): occurrence
in North America, description of a new species,
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Viggiani, G. 1968. Ricerche sugli Hymenoptera Chal-
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Viggiani, G. 1971. Ricerche sugli Hymenoptera Chal-
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dell' armatura genitale esterne maschile dei
Trichogrammatidae. Bollettino del Laboratorio di
Entomologia Agraria 'Filippo Silvestri' di Portici 29:
181-222.
Viggiani, G. 1976a. Ricerche sugli Hymenoptera
Chalcidoidea XLIX. Trichogramma confusum n.
sp. per T. australicum Nagarkatti e Nagaraja
(1968), nee Girault (1912), con note su Tricho-
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gramma heliothidis n. sp. Bollettino del Laboratorio di
Entomologia Agraria 'Filippo Silvestri' di Portici 33:
182-187.
Viggiani, G. 1976b. Ricerche sugli Hymenoptera
Chalcidoidea L. Materiali per una revisione del
genere Oligosita Walk. (Trichogrammatidae). 1. Le
specie australiane descritte da A. A. Girault.
Bollettino del Laboratorio di Entomologia Agraria
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Viggiani, G. 1984. Further contribution to the knowl-
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grammatidae (Hym. Chalcidoidea). Bollettino del
Laboratorio di Entomologia Agraria 'Filippo Silvestri'
di Portici 41: 173-182.
Viggiani, G. 1985. Parassiti oofagi delle cicaline delle
querce: Epoligosita vera Viggiani. Atti Congresso
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Viggiani, G. 1986. Description of a new species of
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Bollettino del Laboratorio di Entomologia Agraria
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Viggiani, G. and S. Laudonia. 1994. Description of
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58(2002): 117-122.

Viggiani, G. and M. Velasquez. 2004. Description of
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Agraria 'Filippo Silvestri' di Portici 59(2003): 87-
91.

Vincent, D. L. and C. Goodpasture. 1986. Three new-
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491-501.

Walker, G. P., I. M. Bayoun, S. V. Triapitsyn, and f. Y.
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(Hemiptera: Cicadellidae) in California. Zootaxa
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Yousuf, M. and Shafee, S. A. 1988. Taxonomy of
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Yoshimoto, C. M. 1976. Pseudoxenufens fbrsythi a new
genus and species of Trichogrammatidae (Hyme-
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The Canadian Entomologist 108: 419-422.

Zucchi, R. A. and R. C. Monteiro. 1997. O genero-
Trichogramma na America do Sul. Chap. 2, Pp. 41-
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gramma e o Controle Biologico Aplicado. FEALQ,
Piracicaba, SP, Brasil. 324 pp.

130

Journal of Hymenoptera Research

Figs 12-19. Antennae. 12, Ceratogramma masneri (9). 13, Ceratogramma jeffersi (9). 14, Ceratogramma jeffersi (,_>).
15, Hydrophylita (9) (arrow to unique spatulate sensilla). 16, Mirufens (9): A - full; B - detail of pedicel showing
crenulate transverse ridges. 17, Mirufens (J) (arrow to small 4th club segment). 18, Pachamama speciosa (9). 19,
Pachamama speciosa (J).

Volume 15, Number 1, 2006

131

Figs 20-29. Antennae. 20, Poropoea (9). 21, Trichogrammatella (9). 22, PJerandrophysalis leventina (9).
23, Pterandrophysalis levantina (<$). 24, Brachyufens osborni (9). 25, Brachyufens osborni | ;). 26, Haeckeliania ^/vr,j/i/
(9). 27, Haeckeliania sperata (<J). 28, Paratrichogramma califbrnica (9). 29, Paratrichogramma California ( ,").

132

Journal of Hymenoptera Research

Figs 30-36. Antennae. 30, Soikiella occidentals (9). 31, Soikiella occidentalis (3). 32, Thanatogramma oweni (9).
33, Thanatogramma oweni (£). 34, Trichogramma pretiosum (9) (arrow to patch of RS). 35, Trichogramma pretiosum
(c?). 36, Trichogrammatoidea bactrae (J).

Volume 15, Number 1, 2006

133

Figs 37-44. Antennae. 37, Trichogrammatomyia (9) (arrow to patch of RS). 38, Trichogrammatomyia ( ,'). 39,
ViggianieUa tropica (9). 40, Xenufens ruskini (9) 41, Xenufens forsythi (9) (lateral, l:l and F2 fused). 42, Xenufens
forsythi (9) (medial, Fl and F2 distinct). 43, Paracentrobia (9): A - full, B - detail of funicle (arrow to F2). 44,
Ittys (9).

134

Journal of Hymenoptera Research

Figs 45-52. Antennae (99). 45, Ittysella lagunera. 46, Aphelinoidea. 47, Brachista fidiae. 48, Ckaetogramma
occidentalis. 49, same (detail of funicle, medial). 50, Lathromeris hesperus (arrow to terminal process). 51,
Nicolavespa theresae (arrow to terminal process). 52, Pintoa nearctica: A - full; B - detail of funicle showing J-
shaped PLS.

Volume 15, Number 1, 2006

135

Figs 53-60. Antennae. 53, Pseuduscana sola (9). 54, same ( ;). 55, Pterygogramma (9). 56, Tumidiclava (9) (arrow to
terminal process). 57, Same (j1). 58, Tumidifemur (9) (note segment asymmetry, especially of C2 and C3). 59, Ufens
(9). 60, same (J1) (arrow to small 4,h club segment).

136

Journal of Hymenoftera Research

Figs 61-68. Antennae. 61, Uscana semifumipennis (9). 62, Xiphogramma fuscum (9). 63, Adelogramma primum (9)
(lateral). 64, same (9) (medial of club). 65, same (^). 66, same (9) (detail of base of club, medial; arrow to very
small CI). 67, Adryas magister (9). 68, Adryas bochica (9). Note CI not visible in lateral views of Adelogramma and
Adryas 9.

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137

Figs 69-76. Antennae. 69, Burksiella spirita (9) (lateral, Fl not visible). 70, same (9) (detail of base of club; note
absence of APB sensilla on CI [compare with Zagella in Fig. 85]). 71, same ( ;) (medial, arrow to Fl). 72,
Chaetostricha (9): A - full; B - detail of funicle (medial, arrow to Fl). 73, Lathromeroidea (9) (Group A). 74,
Lathromeroidea (9) (Group B) (medial, CI visible). 75, same (9) (lateral, CI not visible). 76, same (J) (medial).

138

Journal of Hymenoptera Research

Figs 77-83. Antennae. 77, Lathromeroidea gerriphaga (9) (Group C). 78, Uscanoidea (S). 79, Centrobiopsis odonatae
(9) (medial). 80, same (S) (medial). 81, Zaga sp. 1 (9) (lateral, arrow to CI). 82, same (3). 83, Zaga sp. 2 (9) (lateral,
arrow to CI).

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139

Figs 84-91. Antennae (99). 84, Zagella (medial, arrow to Fl). 85, same (arrow on CI to AI'B sensillum; compare
with Burksiella spirita, Fig. 70). 86, Epoligosita. 87, Megaphragma. 88, Oligosita (collina group). 89, same (detail of
club apex, arrow to clavate PLS). 90, Prestivichia (arrow to Fl). 91, Sinepalpigramma longiciliatum (arrow to Fl).

140

Journal of Hymenoptera Research

Figs 92-98. 92-93, antennae. 92, Doirania elegans (9). 93, Pseudoligosita (9). 94-96, Pterygogramma marquesi
(types). 94, antennal club (9). 95, scutellum (9). 96, 6" genitalia (dorsal; ventral setae are dotted). 97-98,
Viggianiella tropica (9). 97, propodeum and petiolar segment. 98, hind wing.

Volume 15, Number 1, 2006

141

&**■

^J '.

100

-N\\, ■

; ■ '

101

103

■■■-y':r

104

Figs 99-106. Fore wings. 99, Ceratogramma masneri. 100, Ceratogramma jeffersi. 101, Mirufens sp. 1 (arrow to
RSI). 102, Mirufens sp. 2. 103, Pachamama speciosa (arrow to preretinacular lobe). 104, Poropoea. 105,
Pterandrophysalis levantina (9). 106, same (J).

142

Journal of Hymenoptera Research

i-J.

107

•/<

108

■-

...

109

110

111

112

113

114a

JJ^

114

114b

Figs 107-114b. Fore wings. 107, Trichogrammatella. 108, Brachyufens osbomi. 109, Haeckeliania sperata. 110,
Paratrichogramma californica. Ill, Soikiella occidentalis. 112, Thanatogramma oweni. 113, Trichogramma. 114,
Trichogrammatoidea. 114a, Hydrophylita (Hydrophytita). 114b, Hydrophylita (Lutzimicron).

Volume 15, Number 1, 2006

143

116

117

118

<?^

•

119

120

121

122

Figs 115-122. Fore wings. 115, Trichogramtnatomyia. 116, Viggianiella tropica (arrow to stigmal vein). 117,
Xenufens ruskini. 118, Paracentrobia. 119, ftfys (arrow to RSI). 120, lttysella lagunera. 121, Aphelinoidea (Aphelinoidea)
{semifuscipennis Group). 122, Aphelinoidea (Aphelinoidea) (plutella group).

144

Journal of Hymenoptf.ra Research

■

>;-

124

125

126

& i

127

128

129

130

.,-■ ■■■■■■■■■■..■-■■:-.::-W

Figs 123-130. Fore wings. 123, Aphelinoidea (Encyrtogramma). 124, Brachista fidiae. 125, Brachygrammatella. 126,
Chaetogramma occidentalis (arrow to lightly pigmented area at junction of premarginal and marginal veins). 127,
Lathromeris hesperus. 128, Nicolavespa. 129, Pintoa nearctica. 130, Psenduscana sola.

Volume 15, Number 1, 2006

145

132

-. - .

133

135

134

136

. - • - • .-

■

137 138

Figs 131-138. Fore wings. 131, Pterygogramma. 132, Tumidiclava. 133, Tumidifemur sp. 1. 134, Tumidifemur sp. 2.
135, Ufens. 136, L/sram? semifumipennis. 137, Uscanopsis carlyei (type). 138, Xiphogramma fuscum.

146

Journal of Hymenoptera Research

, ■■;

139

140

143

144 b

-^■^*>-N-S

Mm
%

145

Figs 139-146. Fore wings. 139, Adelogramnw primum. 140, Adryas magister (9) (arrow to premarginal vein). 141,
same (J). 142, Burksiella spirita. 143, Chaetostricha. 144, Lathromeroidea (Group A) ('a' to basal convergence of setal
tracks, 'b' to setal line anterior to retinaculum, 'c' to terminus of subcostal vein anterior to base of premarginal
vein). 145, Lathromeroidea exemplum (Group B) (arrow to terminus of subcostal vein anterior to base of
premarginal vein). 146, Lathromeroidea gerriphaga (Group C) (arrow to subcostal vein, confluent with
premarginal vein).

Volume 15, Number 1, 2006

147

/,/

147

148

RS2

r-m

Cu2

~^f

149

151

152

153

154

Figs 147-154. Fore wings. 147, Uscanoidea. 148, Ccntrobiopsis odonatae (with important seta] tracks
characterizing most trichogrammatid wings indicated). 149, Zaga (arrow at RSI). 150, Zagella (arrow at RSI).
151, Epoligosita. 152, Megaphragma. 153, Oligosita. 154, Prestwichia.

148

Journal of Hymenoptera Research

155

156

157

158

Figs 155-162. 155-157, fore wings. 155, Sinepalpigramrna longiciliatum. 156, Doirania elegans. 157, Pseudoligosita.
158-162, fore wing venation (detail). 158, Trichogramma (arrow at campaniform sensilla at apex of premarginal
vein). 159, Aphelinoidea (Aphelinoidea). 160, Tumidifemur. 161, Burksiella spirita (distance between 'a' and 'b' =
distance from base of the marginal vein to the apex of the globose sensilla associated with the stigmal vein
employed for the ratio SV + MV / PM. See Anatomical Structure and Terminology). 162, Zaga.

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149

Figs 163-167. Head capsule. 163, Aphelinoidea (Aphelinoidea) [ See text (Anatomical Structure and Terminologj i
for explanation of acronyms]. 164, Lathromeroidea exemplum (9) (Group B). 165, Lathromeroidea gerriphaga (9)
(Group C). 166, same (c?) (arrow at preocellar pit). 167, Lathromeroidea sp. (Group C), showing detail of preocellar
pit anterior to medial ocellus).

150

Journal of Hymenoptera Research

Figs 168-178. 168-170, mandibles (medial). 168, Oligosita. 169, Trichogramma minutum Riley ('a' to socketed
anterior tooth; 'b' to posterior spine). 170, Hydrophylita (arrow to posterior spine). 171-173, maxillae (posterior).
171, Sinepalpigramma longiciliatum. 172, Oligosita. 173, Mirufens. 174, Hind tibia (posterior), Lathromeroidea
exemplum. 175-178, Fore tibia (dorsal surface above). 175, Aphelinoidea. 176, Chaetostriclw. 177, Zaga. 178, Mirufens.

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Figs 179-182. Mesosoma. 179, Brachyufcm osborni. 180, Lathromeroidea gerriphaga, 181, Lathromeroidea exemplum.
182, Uscanoidea. 183, Lathromeroidea exemplum (sculpturing on midlobe). See text (Anatomical Structure and
Terminology) for explanation of acronyms.

152

Journal of Hymenoptera Research

Figs 184-189. Mesosoma/Metasoma (anterior). 184, Pintoa neurotica (arrow to minute anterior scutellar seta).
185, same (detail of scutellum, arrow to minute anterior seta). 186, OUgosita {collina group, arrow to disk of
propodeum). 187, Pseudoligosita (arrow to disk of propodeum). 188, Paratrichogramma californica (arrow to row of
denticles on first metasomal tergum, SEM). 189, Haeckeliania (row of denticles on first metasomal tergum at
bottom, light microscope).

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153

Figs 190-197. 190-192, mesosoma (lateral). 190, Ceratogramma masneri. 191, Lathromeroidea exemplum. 192,
Oligosita (collina group; ps = mesopleural suture). 193-194, mesophragma. 193, Chaetogramma occidentalis. 194,
Trichogramma. 195-197, full body (lateral). 195, Xiphogramma fuscum (arrow to hypopygium). 196, Lathromeroidea
gerriphaga ('a' to anterolateral bulla on metasomal tergum II; 'b' to hvpopvgium). 197, Psettdoligosita (arrow
to hypopygium).

154

Journal of Hymenoptera Research

Figs 198-199. Male genitalia of Trichogramma sp. (labeled). 198, dorsal
Structure and Terminology) for explanation of acronyms.

199, ventral. See text (Anatomical

Volume 15, Number 1, 2006

155

Figs 200-207. Male genitalia. 200, Ceratogramma masneri (dorsal). 201, same (ventral; 'a' to paramere, V to
volsella, 'c' to intervolsellar process, 'd' to aedeagus). 202, Ceratogramma jeffersi (ventral). 203, Hydropkylita
(Lutzimicron) (ventral). 204, same (detail of apex). 205, Mirufens (dorsal; 'a' to paramere, V to volsella, V to
aedeagus. 206, same (ventral). 207, Pterandrophysalis levantina (ventral).

156

Journal of Hymenoptera Research

Figs 208-214. Male genitalia. 208, Trichogrammatella (ventral). 209, Haeckeliania sperata (dorsal). 210,
Paratrichogramma californica (dorsal; arrow to basal fusion of aedeagal apodemes). 211, same (ventral). 212,
Soikiella occidentalis (dorsal). 213, same (posterodorsal; arrow to aedeagal apodemes). 214, same (ventral; arrow to
paramere - volsella hidden).

Volume 15, Number 1, 2006

157

Figs 215-223. Male genitalia. 215, Thanatogramma oweni (dorsal; aedeagus removed). 216, same (aedeagus
drawn to same scale as Figs 215 and 217). 217, same as 215 (ventral). 21S, Trichogramma sp. (dorsal). 219,
Trichogramma sp. (ventral). 220, Trichogrammatoidea bactrae (dorsal). 221, same (ventral). 222, Trichogrammatomyia
(ventral). 223, Xenufens ruskini (ventral).

158

Journal of Hymenoptera Research

Figs 224-230. Male genitalia. 224, Paracentrobia (arrow to a strapshaped appendage of last metasomal
sternum). 225, Ittys ('a' to a platiform appendage of last metasomal sternum, 'b' to sickle shaped volsellus). 226,
Ittysella lagunera (arrow to sickle shaped volsella). 227, Aphelinoidea (plutella group) (ventral; arrow to aedeagal
apodeme). 228, same (ventral; 'a' to ventral seta, 'b' to volsella). 229, Chaetogramma occidentalis (dorsal). 230, same
(ventral; 'a' to ventral seta, 'b' to volsella).

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159

Figs 231-238. Male genitalia. 231, Lathromeris hesperus (ventral). 232, Nicolavespa theresae (dorsal). 233, same
(ventral). 234, Pseuduscana sola (dorsal). 235, same (ventral; 'a' to paramere, V to vulsella). 236, Tumidiclava
(dorsal; arrow to apparent demarcation line between capsule and aedeagus). 237, same (ventral; demarcation
line on dorsal side not occurring ventrally). 238, Ufens principalis (dorsal).

160

Journal of Hymenoptera Research

Figs 239-248. Male genitalia. 239, Xiphogramma fuscum (dorsal). 240, same (ventral). 241, Adelogramma primum
(dorsal). 242, Burksiella spirita (dorsal). 243, same (ventral). 244, Chaetostricha (dorsal). 245, same (ventral). 246,
Lathromeroidea (Group C) (dorsal). 247, same (ventral; base distorted). 248, Latliromeroidea gerriphaga (Group
C) (dorsal).

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161

Figs 249-257. Male genitalia. 249, Lathromeroidea (Group B) (ventral; arrow to ventral seta). 250, Uscanoidea
(ventral). 251, Centrobiopsis odonatae (ventral; note longitudinal furrow). 252, Zaga (dorsal). 253, same (ventral).
254, Zagella (dorsal; North American species). 255, same (ventral; arrow to ventral seta). 256, Zagella (ventral;
South American species). 257, Same (detail of apex; arrow to paramere or volsella).

162

Journal of Hymenoptera Research

Figs 258-265. 258-259, male genitalia (Oligositini). 258, Oligosila (arrow to anterodorsl aperture). 259,
Pseudoligosita (ventral; 'a' to anterodorsal aperture [note posteriorly directed apodemes below]; 'b' to apex of
genitalia). 260-261, last sternal area in male. 260, Zagella. 261, Lathmmeroidea gerriphaga (Group C). 252-264,
dorsal metasomal modifications in Lathromeris. 262, Lathromeris sp. 1. 263, Lathromeris hesperus. 264, Lathromeris
sp. 265, Brachista fidiae (9) (arrow to strut-like apodeme anterior to ovipositor)

Volume 15, Number 1, 2006

163

tf 'WA

wMt

\ '

266

•>.•;

267

272

268

/■

273

271

274

Figs 266-274. Burksiella dianae. 266, habitus (9). 267, antenna (9) (arrow to suture between I! and F2). 268,
same (J). 269, maxillary palp. 270, fore wing. 271, midlobe of mesoscutum showing sculpturing. 272, female
metasoma. 273, male metasoma. 274, male genitalia (arrow to cuticular flange at base).

J. HYM. RES.
Vol. 15(1), 2006, pp. 164-170

Nomenclatural Changes in Old World Crabronidae (Hymenoptera),
with Taxonomic Comments and New Distribution Records

WOJCIECH J. PULAWSKI

California Academy of Sciences, Golden Gate Park, San Francisco, California 94118, USA;

email: wpvilawski@calacademy.org

Abstract. — The following nominal species are newly synonymized (the valid names are listed
last): Cerceris insularis F. Smith, 1856 = Cerceris armaria (Linnaeus, 1758); Oxybelus rufipes
Taschenberg, 1880 = Oxybelus lamellatus Olivier, 1811; Tachytes astatiformis Tsuneki, 1963 =
Tachytes pygmaeus Kohl, 1888; Tachytes decorsei Berland, 1942, and falciger Arnold, 1951 = Tachytes
chudeaui Berland, 1942; Tachytes diversicomis Turner, 1918, niger Berland, 1942, senegalensis Berland,
1942, tassilicus Pulawski, 1962, and hengchunensis Tsuneki, 1967 = Tachytes xenoferus Rohwer, 1911;
Tachytes eurous Pulawski, 1962 = Tachytes aeneus Saunders, 1910; and Tachytes rostratus Berland,
1942 = Tachytes basilicas Guerin-Meneville, 1844. A new name, Tachytes dogon, is proposed for
Tachytes rufipes Berland, 1942, a junior primary homonym of Tachytes rufipes Aichinger, 1870.

My fieldwork in various parts of the Old
World, as well as visits to museums,
revealed previously unnoticed synony-
mies, which are discussed below. The
relevant types have been examined, except
for that of Oxybelus lamellatus Olivier which
is no longer in existence, for Cerceris
arenaria Linnaeus whose identity has been
firmly established (Richards 1935, Day,
1979), and for Tachytes basilicus Guerin-
Meneville whose identity was confirmed
by Pulawski (1962). The abbreviations used
in the text for the institutional or personal
collections that house these types or other
specimens discussed are:

BMNH: The Natural History Museum,
formerly British Museum (Nat-
ural History), London, United
Kingdom

CAS: California Academy of Sciences,

San Francisco, California, USA

CSE: Christian Schmid-Egger, Herr-

sching-Breitbrunn, Germany
(personal collection)

GENOVA: Museo Civico di Storia Natur-
ale "Giacomo Doria", Genova,
Italy

HALLE: Institut fur Zoologie, Martin-
Luther Universitat Halle Wit-
tenberg, Halle, Germany

KOBE: Entomological Laboratory, Fac-
ulty of Agriculture, Kobe Uni-
versity, Rokko, Kobe, Japan

MNHN: Museum National d'Histoire
Naturelle, Paris, France

NHMW: Naturhistorisches Museum,
Wien, Austria

OXUM: Oxford University Museum of
Natural History, Oxford, Unit-
ed Kingdom

USNM: U.S. National Museum of Nat-
ural History, Smithsonian In-
stitution, Washington, D.C.,
USA

Cerceris arenaria

Sphex arenaria Linnaeus, 1758:571. Holotype: 9,
Sweden: no specific locality (Linnean Society,
London). - Richards, 1935:169 (study of
holotype); Day, 1979:50 (study of holotype).

Cerceris insularis F. Smith, 1856:444, 9. Holotype
or syntypes: 9, Italy: Sicily: no specific locality
(BMNH), examined. New synonym. - Schlet-
terer, 1887:441 (as tentative synonym of
Cerceris ferreri); Dalla Torre, 1897:460 (listed

Volume 15, Number 1, 2006

165

as synonym of Cerceris ferreri). - As Cerceris
flavilabris insularis: Bohart and Menke,
1976:581 (listed); Pagliano, 1990:97 (in catalog
of Italian Sphecidae).

The only specimen of Smith's insularis in
the BMNH is a typical female of the
common Palearctic species armaria, with
a distinctive clypeal lamella.

Oxybelus lamellatus Olivier

Oxybelus lamellatus Olivier, 1811:595, sex not
indicated. Syntypes: Egypt: no specific local-
ity; and Iraq: Baghdad area (destroyed).

Oxybelus rufipes Taschenberg, 1880:781, 9, J.
Lectotype: 9, Ethiopia: no specific locality
(HALLE), present designation. New syno-
nym.

I follow de Beaumont (1950:413) in his
interpretation of Oxybelus lainellatus. The
species is characterized by the following
combination: mucro reddish brown, folia-
ceous, emarginate apically; postscutellum
yellow between lamellae (except in many
males), lamella divided apically; sternum II
coarsely punctate except punctures con-
trastingly fine laterally; antennal flagellum
and femora red; pygidial plate of female
unusually broad; and male terga III-VI
with conspicuous lateral spines. All these
characters are found in the syntypes of
Oxybelus rufipes.

Tachytes aenens Saunders

Tachytes aeneus Saunders, 1910:522, £ (as aenea,
incorrect original termination). Syntypes: S,
Algeria: Biskra (OXUM), one syntype exam-
ined before 1962. - Morice, 1911:99 (Algeria:
Biskra); de Beaumont, 1955:172 (Morocco:
Marrakech); Pulawski, 1962:421 (in revision
of Palearctic Tachytes); R. Bohart and Menke,
1976:263 (listed).

Tachytes eurous Pulawski, 1962:423, 9. Holotype:
9, Israel: Eyn Aosb, now Hatzeva (originally
H. Bytinski-Salz collection, now Tel Aviv
University). New synonym. - de Beaumont,
Bytinski-Salz and Pulawski, 1973:7 (Israel); R.
Bohart and Menke, 1976:265 (listed).

I described Tachytes eurous from a single
female from Israel, because I suspected it

may not be conspecific with the western
North African aeneus, known only from the
male. A recent study of topotypical females
and males from both Morocco and Israel
convinced me that only one species is
involved. These two names are therefore
synonyms.

Tachytes aeneus has a unique propodeal
dorsum that has a triangular, ridged,
glabrous area extending from base to apex
(orientation of ridges varies from trans-
verse to longitudinal). Also, the erect setae
are all or largely absent on the scape, but
present on tergum I. The female clypeus,
with its free margin emarginate mesally
and sharply pointed lip corner, is also
distinctive. For additional characters, see
Pulawski (1962:420).

Material examined. — ISRAEL: En Shahak in
En Yahav Makhteshim Reserve at 30 42.8' N
35°11.1'E (1 9, 1 J, CAS; 1 3, CSE), Iddan in
Arava Valley at 3048.9'N 3516.8'E (1 9, 2 6\
CSE). MOROCCO: Marrakech (1 9, 1 ?, CAS).

Tachytes basilicus (Guerin-Meneville)

Lyrops basilicus Guerin-Meneville, 1844:440, 9.
Holotype: E, Senegal: no specific locality
(GENOVA). - As Tachytes basilicus: F. Smith,
1856:300 (new combination, listed, spelled
basilica); Pulawski, 1962:416 (full bibliogra-
phy, synonymy, revision).

Tachytes rostratus Berland, 1942:6, j (as wstrata,
incorrect original termination). Holotype: ',
Chad: Moyen Chari: Fort Archambault, now
Sarh (MNHN), examined. New synonym.

The holotype of rostratus, a badly worn
specimen with gastral pubescence matted
by moisture, has all the distinctive char-
acters of basilicus described by Pulawski
(1962). I could not find any noteworthy
differences when comparing it to speci-
mens of basilicus.

The most important recognition features
of basilicus are (Pulawski 1962): mouthparts
elongate (in particular, galea markedly
longer than scape and without transverse
suture); clypeal free margin only slightly
concave between lobe and eye orbit;
hindfemur without apical lobe; pilosity

166

Journal of Hymenoptera Research

concealing integument at scutal forecor-
ners; and gaster red at least basally, with
appressed golden vestiture. In the female,
the setae of the pygidial plate do not
conceal the integument, and in the male
the apical margin of sternum VIII is only
slightly emarginate, almost entire.

Tachytes chudeaui Berland

Tachytes chudeaui Berland, 1942:3, 9 (as Chudeaui,
incorrect original capitalization). Holotype: 9,
Mali: Niafunke near Toumbouctou (MNHN),
examined. - R. Bohart and Menke, 1976:264
(listed).

Tachytes decorsei Berland, 1942:7, J (as Decorsei,
incorrect original capitalization). Holotypeo,
Mali: Douentza (MNHN), examined. New
synonym. - R. Bohart and Menke, 1976:264
(listed).

Tachytes falciger Arnold, 1951:152, j (as falcigera,
incorrect original termination). Holotype: $,
Mauritania: Aleg (BMNH), examined. New
synonym. - R. Bohart and Menke, 1976:265
(listed).

The species is a typical member of the
obsoletus group as defined by Pulawski
(1962). It can be recognized by the follow-
ing characters: postocellar area narrow
(least interocular distance equal 1.1 X
length of flagellomere I in the female and
0.9-1.4 X in male), with erect setae and
most punctures less than one diameter
apart, apical lobe of hindfemur relatively
large, as in archaeophilus Pulawski (see Figs
71 and 72 in Pulawski, 1962), hindfemoral
venter without erect setae, tergum I with
erect setae (only laterally in female and
small males), femora and tibiae all black.
Female: clypeal bevel rudimentary, terga I
and II red (remainder black), tergum V
with silvery pubescence laterally. Male:
ventral margin of flagellomeres VIII and
IX expanded (in many specimens also that
of flagellomere X), gaster all black.

The species was previously known from
Mauritania and Mali, but it also occurs in
Burkina Faso and Niger.

Material examined (all CAS). —BURKINA
FASO: Pala (1 S). NIGER: Agadez Region:

5 km N Agadez at 17 01.2'N 800.7'E (1 9, 16 $),
30 km S Agadez at 16 39.0'N 7 56.9'E (1 3)-
Diffa Region: 3 km N Diffa at 13 21.3'N
12 36.7'E (1 S), 8 km N Diffa at 13 24.1 'N

12 36.2'E (3 9, 1 J), 54 km NE Diffa at

13 42.3'N 12 55.8'E (3 1), 87 km NE Diffa at

14 02.9'N 12 58.5'E (1 9), 14 km W Diffa at
13T5.8'N 12 29.0'E (2 $), 34 km SW Nguigmi at
13 58.8'N 12 58.2'E (1 9), 42 km SW Nguigmi at
13 54.5'N 12 56.5'E (4 9). Dosso Region: 13 km
S Dosso at 1256.6'N 311.0'E (3 9) 15 km N
Gaya at 11 59.6'N 3 32.2'E (1 J). Maradi
Region: 15 km NNW Maradi at 13 37.9'N
7 03.0'E (3 S), 17 km NNW Maradi at
13 38.7'N 7 026'E (1 J). Tahoua Region: Ta-
houa at 14 53.5'N 5 16.6'E (3 9). Tillaberi
Region: 11 km N Ayorou at 14 49.3'N 0 52.2'E
(2 $), 8 km SE Kollo at 13 16.4'N 2 22.0'E (1 9),
Malale 10 km E Niamey at 13 27.1 'N 2 10.4'E (1
9), 21 km N Niamey at 13 33.2'N 2 21.5'E (3 9),
63 km NW Niamey at 13 53.4'N 1 35.2'E (1 S),
82 km ESE Tera at 13 51.1'N 1 31.3'E (2 $),

15 km NW Tillaberi at 14 17.3'N 1 20.5'E (1 9,
1 J). Zinder Region: 21 km W Goure at
13 51.2'N 10 07.8'E (1 9), 3 km S Takieta at
13 39.6'N 8 30.7'E (1 $), 6 km S Takieta at

13 37.1'N 8 30.6'E (1 $), 45 km S Tanout at

14 37.4 'N 8 44.3 'E (1 J).

Tachytes dogoti Pulawski, new name

Tachytes chudeaui var. rufipes Berland, 1942, 9, $
(9 = Tachytes saharicus), junior primary
homonym of Tachytes rufipes Aichinger,
1870, which is a junior synonym of Tachy-
sphex brullii (F. Smith, 1856). Lectotype: $,
Mali: Douentza (MNHN), designated by
Pulawski, 1962:385, reexamined in 2005. -
As Tachytes rufipes: Pulawski, 1962:385 (new
status).

Named after the Dogon people of Mali.

This species can be recognized by the
following characteristics. Galea shorter
than scape, female clypeus noncarinate
and apical depression of sternum II im-
punctate, male forecoxa and foretrochanter
not modified. The following are red: scape,
flagellum partly, femora, tibiae, and tarsi;
wings yellow, infumate along distal mar-
gin (infumate portion wears off in old
specimens); gaster at least partly red.
Hindfemur without apical lobe at apicov-

Volume 15, Number 1, 2006

167

entral angle of outer side (see Figure 731 in
Bohart and Menke, 1976, for lobe present).
In female, punctures of pygidial plate
markedly more spaced than average in
genus, especially anterolaterally (integu-
ment easily visible between the setae
except posteriorly). In male, width of
postocellar area (= least interocular dis-
tance) about equal to midocellar width;
midbasitarsal venter evenly curved except
somewhat expanded apically, without
spines; apex of sternum VIII only slightly
emarginate (markedly less than average for
the genus). Hindfemoral venter with no
erect setae in female and many males, but
a few suberect setae present on basal half
in some males (setal length no greater than
midocellar width).

Material examined (all CAS): MALI: 10 km E
Mopti (4 9, 10 $). NIGER: Diffa Region: 42 km
SW Nguigmi at 13 54.5'N 12 56.5'E (1 $).
Tillaberi Region: 11 km N Ayorou at
14°49.3'N 0 52.2'E (1 J), 82 km ESE Tera at
13°51.1'N 1 31.3'E (2 J). Zinder Region: 19 km
E Goure at 13 52.6'N 10 24.9'E (1 $), 27 km W
Guidiguir at 13 40.9'N 9 39.1'E (2 £), 29 km
NW Magaria at 13 09.1 'N 8 41.3'E (1 $), 3 km S
Takieta at 13 39.6'N 8 30.7'E (6 $), 45 km S
Tanout at 14 37.4'N 8 44.3'E (1 $).

Tachytes pygmaeits Kohl

Tachytes pygmaea Kohl, 1888:134, &<?(<? =
Tachytes argyreus F. Smith), incorrect original
termination. Lectotype: 9, Egypt: no specific
locality (NHMW), designated by Pulawski,
1962:465.

Tachytes astaiiformis Tsuneki, 1963:6, ._>• Holo-
type: o, Thailand: Chiangmai (KOBE), exam-
ined. New synonym. - R. Bohart and Menke,
1976:236 (listed).

Tachytes pygmaeus is easily recognized by
its glabrous propodeal dorsum combined
with an emarginate posterior mandibular
margin and the propodeal spiracle sepa-
rated from the postnotum by less than its
own length. Tachytes dichrous F. Smith is
the only other member of the genus with
an entirely asetose propodeal dorsum.
Unlike pygmaeus, the mandibular posterior

margin of dichrous is not emarginate, and
the propodeal spiracle is separated from
the postnotum by more than its own
length, among other characters.

Pulawski (1962:465) gave Egypt, Moroc-
co, and Sudan as the geographic distribu-
tion of pygmaeus, but Bohart and Menke
(1976:266) correctly listed if from all of
Africa, India, and Sri Lanka. The new
synonymy extends its range to Thailand.

Tachytes xenoferus Rohwer

Tachytes xenoferus Rohwer, 1911:581, 9, q\
Holotype: S, India: Gujarat: Deesa (USNM),
examined.

Tachytes diversicornis R. Turner, 1918:94, J, 9.
Lectotype: $, Pakistan: Karachi (BMNH),
designated by Pulawski, 1975a:316, exam-
ined. New synonym.

Tachytes Chudeaui var. niger Berland, 1942:4, 9
(as nigra, incorrect original termination).
Holotype: 9, (Mali?): Middle Niger basin:
Siganara (MNHN), reexamined in 2005. New
synonym.- As Tachytes niger: Pulawski,
1962:402 (new status). - As Tachytes chudeaui
niger: Bohart and Menke, 1976:264 (new
status, listed).

Tachytes senegalensis Berland, 1942:9, > Holo-
type:^, Mali: Kayes (MNHN), examined in
2005. New synonym.

Tachytes tassilicus Pulawski, 1962:401, J. Holo-
type: Algeria: Djanet (MNHN), reexamined
in 1991. New synonym.

Tachytes liengchunensis Tsuneki, 1967:47, 9, j1.
Holotype: E, Taiwan: Pingtung County:
Hengchun (originally K. Tsuneki coll., now
USNM), examined in 2005. New synonym. -
Haneda, 1971:30 (Taiwan); Tsuneki, 1971:8
(Taiwan); Haneda, 1972:4 (Taiwan; as feng-
chunensis); Murota, 1973:118 (Taiwan); R.
Bohart and Menke, 1976:265 (listed); Tsuneki,
1977:269 (Taiwan); Nuhn and Menke, 1994:25
(holotype transferred to USNM); Porter,
Stange, and Wang, 1999:8 (in checklist of
Sphecidae of Taiwan).

Tachi/tes xenoferus is a member of the
maculicornis group as defined by Pulawski,
1962:390. It is characterized by an all black
gaster and the absence of erect setae on the
midfemoral venter (erect setae may be
present or absent on sternum II). Other

168

Journal of Hymenoptera Research

characters include: mouthparts short
(galea shorter than scape), propodeal
dorsum and side not ridged, female
clypeus not carinate, apical depression of
sternum II impunctate, and pygidial plate
somewhat acute apically, male forecoxa
and foretrochanter not modified. Both
sexes can be recognized from simi-
lar species by midtarsomere III slight-
ly shorter than II (rather than equal in
length).

The male of Tachytes xenoferus is charac-
terized by the midbasitarsus expanded
apically, midtarsomere II not modified,
sternum VIII with usual, short setae, most
or all of the clypeal bevel densely punctate
(punctures less than one diameter apart or
nearly so), and in many specimens middle
flagellomeres yellowish to a various de-
gree. These characteristics are shared by
the male of diversicornis, which clearly is
a junior synonym. The hindfemur is
minimally concave basoventrally in many
specimens (including the holotypes of
xenoferus and diversicornis). Also, flagello-
meres IX and X are only minimally
expanded apicoventrally (in the holotype
of xenoferus, a stylopized specimen, only
flagellomeres I and II are preserved on one
antenna, and I- VI on the other). The other
species in which the male has an apically
expanded apex of midbasitarsus differ
from xenoferus as follows (in addition to
the midtarsomeres II and III). In argenteus
Gussakovskij, the clypeal bevel has only
a few, sparse punctures. In flagellarius
Nurse, flagellomeres IV-X are markedly
expanded ventrally and the hindfemur is
markedly concave basoventrally. In macu-
licornis E. Saunders, flagellomeres III-VII
are roundly expanded ventrally, the hind-
femur is markedly concave basoventrally,
and midtarsomere II is asymmetrical, with
apex expanded anteroventrally. In sacricola
Pulawski, the hindfemur is markedly con-
cave basoventrally and midtarsomere II is
asymmetrical, with apex expanded ante-
roventrally. Finally, in trichopygus Pu-
lawski (female unknown), setae of sternum

VIII are conspicuously elongate (apical
setae 3 X midocellar diameter).

I described Tachytes tassilicus from a sin-
gle male and I thought that it differed from
diversicornis (i. e., xenoferus) in having the
hindfemoral venter not concave basally
and the thoracic setae golden (contrasting
with gastral setae). A study of additional
material convinced me that these differ-
ences do not stand scrutiny. In fact, the
hindfemoral venter varies from slightly
concave to entire, and the thoracic vestiture
may be silvery in specimens with noncon-
cave hindfemoral venter, and golden in
specimens with a slightly concave venter.
Clearly, these two names are synonyms (as
well as synonyms of xenoferus). The holo-
type of Tachytes senegalensis is identical to
tassilicus and is therefore another synonym
of xenoferus.

Specimens from Niger vary markedly in
color. In most, the femora and the tibiae are
black. In several females, however, part of
the forefemur and the mid- and hindfe-
mora are red, and in some all the femora
are red. In several males, all the tibiae and
the hindfemur are red (the hindfemur may
be black basally), and in some also a part of
the midfemur is red.

I thought (Pulawski, 1962:403) that Ta-
chytes niger Berland was an invalid junior
homonym of Tachytes niger Vander Linden,
1829. This opinion was incorrect, as Vander
Linden (1829) did not describe the species,
but transferred to Tacliytes the species
Sphex niger Fabricius, 1775 (which is cur-
rently placed in Liris, see International
Commission on Zoological Nomenclature,
1973, Opinion 997).

Tachytes xenoferus was described from
the Gujarat State of India, while Pulawski
(1962:401) listed it (as diversicornis) from
Pakistan, Israel, Egypt, and Sudan, and
Bohart and Menke (1976:264) added
Ghana, Ethiopia, and Mali. I collected the
species in Burkina Faso, Niger, and Oman,
and I examined specimens from Saudi
Arabia, India, Sri Lanka, Thailand, and
Taiwan.

Volume 15, Number 1, 2006

169

Material examined (all CAS): BURKINA
FASO: 36 km NE Bobo Dioulasso at 11 23.3' N
4°04.3'W (2 9, IS), 71 km NNW Bobo Dioulasso
at 1142.0'N 4 31.4'W (2 J), 28 km NE Dedou-
gou at 12°35.5'N 3°15.6'W (2 9), 38 km SSW
Dedougou at 12 10.5'N 3 36.5'W (5 $), 10 km E
Dori 14 00.5'N 0 03.2'E (1 $), 39 km E Dori
13°58.1'N 0 17.5'E (1 3), 4 km NE Dori at
14°03.8'N 0 03.1 'E (2 9, 3 $), 15 km S Gorom
Gorom at 14 21.4'N 0°07.9'E (2 9, 15 $), 36 km E
Koudougou at 12 12.0'N 2 01.6'W (1 9), 69 km
W Koudougou at 1214.8'N 2 57.7'W (1 9),
17 km S Koupela at 12°02.2'N 0 21.8'E (1 9, 1 3),
80 km S Ouagadougou at 11°40.9'N T 14.2'W (1
3), 13 km NE Ouahigouya at 13 38.9'N
2°19.6'W (1 3), 4 km NW Ouahigouya at
13 37.0'N 2 27.6'W (2 9, 12 3), 15 km SSE
Ouahigouya at 13°27.0'N 2'22.9'W (1 3),
52 km SSE Ouahigouya at 13 07.3'N 2 20.8'W
(1 3), 2 km W P6, 1111.0'N 1 09.5'W (3 6*)-
INDIA: Pondichery Territory: Karikal (4 3).
Tamil Nadu: Coimbatore (1 9). NIGER: Agadez
Region: 0.5 km SE Aderbissinat at 1536.9'N
7°54.0 E (1 9), 5 km N Agadez at 17'01.2'N
8°00.7'E (1 9, 6 3). Dosso Region: 13 km S
Dosso at 1256.6'N 311.0'E (1 $), 39 km S
Dosso 12°40.1'N 3T0.6'E (6 3), 15 km N Gaya at
11°59.6'N 332.2'E (1 9, 1 J). Maradi Region:
23 km NNW Maradi at 13°42.3'N 7 01.4'E (1 3).
Niamey Region: 8 km NW Niamey at 13°35'8N
1°59.9'E (2 9, 1 3). Tillaberi Region: 13 km N
Ayorou at 14 50.1 'N 0'52.4'E (1 9, 2 £), 2 km SE
Kollo at 13T9.6'N 2T9.9'E (1 9, 1 ?), Malale
10 km E Niamey at 13 27.1 'N 2 10.4'E (1 9, $),
13 km N Niamey at 13°32.6'N 2T6.4'E (1 9),
15 km NW Tillaberi at 14T7.3'N 1 20.5'E (1 3),
30 km SSW Torodi at 12 49.5'N (1 9). Zinder
Region: Bosotchouwa 20 km SW Takieta at
13 30.1 'N 8 31.9'E (1 $), 49 km SW Dengas at
12°52.8'N 8 58.6'E (1 9), 18 km N Goure at
14°08.7'N 10 11.6'E (1 9, 1 3), 21 km W Goure at
13°51.2'N 10 07.8'E (1 9), 23 km NNW Maradi
at 13 42.3'N 7 01.4'E (1 9), 3 km S Takieta at
13°39.6'N 8 30.7'E (3 9, 1 $), 6 km S Takieta at
13°37.1'N 8 30.6'E (1 9), 31 km NW Tanout at
15°05.6'N 8 36.1 'E (1 3), 11 km S Tanout at
14°52.6'N 8 52.3'E (1 9), 18 km S Tanout at
14°48.3'N 8 51.9'E (1 3), 37 km S Tanout .it
14°38.2'N 8 42.6'E (2 3), 45 km S Tanout at
14°37.4'N 8 44.3'E (1 9), 55 km S Tanout
14°31.2'N 8 44.3'E (1 3), 44 km N Zinder
14°12.9'N 8 49.3'E (1 9, 1 3), 52 km N Zinder
at 14°17.2'N 8°46.9'E (1 9), 37 km S Zinder at

14°13.3'N 9°00.5'E (4 9, 2 3), 45 km S Zinder at
13°27.3'N 9°00.5'E (1 3). OMAN: Wadi Ghul
near Nizwa at 22 53.0'N 5731.2'E (1 9). SAUDI
ARABIA: El Riyadh (2 9, 5 i), Haddat Ash
Sham (1 9). SRI LANKA: Colombo District:
Ratmalana (1 9). Mannar District: Kokmotte
Bungalow 0.5 mi. NE Wilpattu (1 ;). TAIWAN:
Pingtung Shih (county): Kentin (1 j), Manchou
(1 9, 1 J). Taitung Shih: Tulan (1 9). Shih
unknown: Anping (1 j), Taihorin (1 9). THAI-
LAND: Songkhla Province: Hat Yai (1 9).

ACKNOWLEDGMENTS

I sincerely thank the museum curators who sent
material for study: M. Dorn (Halle, Germany),
Larraine Ficken (London, United Kingdom), Ted
Schulz and the late Karl V. Krombein (Washington,
D.C., USA), Tikahiko Naiko (Kobe, Japan), and Claire
Villemant (Paris, France). Jere S. Schweikert and
Arnold S. Menke kindly reviewed earlier drafts of
the manuscript.

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J. HYM. RES.
Vol. 15(1), 2006, pp. 171-176

Plants Whose Flowers are Utilized by Adults of Pepsis grossa Fabricius
(Hymenoptera: Pompilidae) as a Source of Nectar

Fred Punzo

Department of Biology, Box 5F, University of Tampa, 401 W. Kennedy Blvd., Tampa, Florida

33606-1490, USA; email: fpunzo@ut.edu

Abstract. — The spider wasp, Pepsis grossa Fabricius is a common species of wasp found in the
Chihuahuan Desert. This study was conducted in Big Bend Ranch State Park (Brewster Co., Texas)
and reports on flowers of various plant species used by adults of this wasp as a source of nectar.
Although nectar was obtained from the flowers from a total of 19 plant species, four of those species
(milkweed: Ascepias texana and A. sperryi; Mexican Buckeye: Ungadia specisosa; Honey Mesquite:
Prosopis glandulosa ) accounted for 73.6% of all plants utilized. The flowers from these plants were
all characterized by short corolla tube lengths (<3.5 mm). At this site, P. grossa exhibited a narrow
trophic niche breadth (Levin's index: B = 0.2816). Other common species of plants whose flowers
were not visited by these wasps are also identified.

Tarantula hawk wasps of the genus
Pepsis (Hymenoptera: Pompilidae: Pepsi-
nae: Pepsini) are large, long-legged wasps,
and are conspicuous components of the
arthropod fauna of desert regions of the
southwestern United States and Mexico
(Punzo 1994a, 2000). The genus Pepsis is
found in North and South America, and
the West Indies, ranging from Utah (lati-
tude 42 N) to Argentina (45 S) (Hurd 1952,
Vardy 2002). Although females of these
wasps typically hunt and paralyze large
spiders as a source of food for their
carnivorous larvae (Petrunkevitch 1926,
Schmidt 2004), the adults are nectivorous
(Williams 1956, Punzo 1994b).

Although there have been some studies
on the ecology (Cazier and Mortenson
1964, Field 1992, Punzo 2005), territoriality
(Alcock 1979, 2000), hunting behavior
(Petrunkevitch 1926, Cazier and Mortenson
1964, Punzo and Garman 1989, Punzo
1994b), neurochemistry (Punzo 1990,
1991), venom chemistry (Schmidt 1990,
2004), morphology of venom glands (Scho-
eters et al. 1997), and physiology (Punzo
1990) of Pepisis wasps, there is a paucity of

information available on the natural histo-
ry of many species (Punzo 2005).

These wasps are known to obtain nectar
from the flowers of a variety of plants
(Evans and West-Eberhard 1970, Punzo
2000). However, little information exists
on the specific range of plants utilized by
any Pepisis wasp (Punzo 2005). In desert
regions, where ambient temperatures may
exceed 43 C during late spring and sum-
mer months, female wasps often fly over
considerable distances during daylight
hours, searching for a suitable host (Vardy
2002, Punzo 2005). Male wasps engage in
energetically costly activities as well, in-
cluding an aggressive aerial defense of
territories against intruders (Alcock and
Bailey 1997). Insect flight is energetically
costly under any circumstance. Thus, it is
essential that adult wasps obtain required
nutrients in order to survive and repro-
duce. In addition, because any specific type
of plant tissue or nectar may lack some
essential dietary requirement, it is only
through careful selection of particular
plants that an animal can obtain a balanced
diet.

172

Journal of Hymenoptera Research

The purpose of this study was to identify
the species of plants (flowers) visited by
adults of Pepisis grossa (Fabricius 1798),
formerly described as Pepsis formosa for-
mosa (Say), in order to obtain nectar. The
study was conducted in Big Bend Ranch
State Park located in Trans Pecos Texas.

DESCRIPTION OF GENERAL
STUDY AREA

Big Bend Ranch State Park (BBRSP,
Presidio Co., Texas) lies within the north-
ern region of the Chihuahuan Desert. It is
bordered to the west by the Rio Grande
River (RGR), and climatic conditions range
from semi-arid to arid (Parent 1996, Punzo
2000). Mean monthly air temperatures
range from 5.1 C in January to 33.4 C in
July, with low and high temperatures of
-10.7 and 45.6 C, respectively (U.S. Dept.
of Interior, 2002). Annual rainfall is typi-
cally between 14.6-29.7 cm, depending on
location and altitude, with 70-80% occur-
ring from May through October (Medellin-
Leal 1992). A wide range of topographic
diversity exists within the Park, including
igneous rocks, gypsum formations, lime-
stone deposits that provide a variety of
substrates including alluvial fans, moutain
ridges, canyons, saline playas, gypsum
flats, siliceous and gypsum dunes, fine-
textured basins, and freshwater seeps and
springs (Milford 1991), all supporting a di-
verse plant fauna with distinct vegetative
zones (Powell 1988).

MATERIALS AND METHODS

I conducted field studies within the
BBRSP in 2003, from late March through
September, when plants of this region have
well developed flowers. The study site was
an area located within a 3.0 km radius of
Grassy Banks (29 17'30" N, 103 55 '04" W;
elevation: 814 m) which is located directly
off State Road 170, 6.76 km NW of Lajitas,
Texas. The western edge of the site is
bordered by the RGR. The soils along the
canyon floor are a mixture of sand, gravel
and adobe, and support a predominantly

sotol-lechuguilla plant community. The
dominant vegetation includes lechuguilla
(Agave lechuguilla), smooth sotol (Dasylirion
leiophyllum), ocotillo (Fouquieria splendens),
mesquites (Prosopis spp.), purple sage
(Leucophyllum frutescens), creosote (Larrea
divaricata), false agave (Hechtia texensis),
tarbrush (Flourensia cemua), catclaw acacia
(Acacia berlandieri), prickly pear cacti
(Opuutia spp.), yuccas (Yucca spp.), and
scattered clumps of milkweed (Ascepias
spcrryi) and grasses, including chino
gramma (Bouteloua breviseta), fluffgrass
(Erioneuron pulchellum), and beargrass (No-
lina erumpens). Numerous large rocks and
boulders are scattered along the canyon
floor, and the canyon walls are comprised
mainly of sandstone, limestone, and igne-
ous rock.

Pepsis grossa (Fabricius) is the most
common Pepsis wasp at this location
(Punzo 2000). Males can be observed in
flight either moving between flowers or
defending territories, feeding at flowers,
or resting on various bushes and plants.
Females are readily visible, flying in
search of hosts or food plants, or walking
rapidly over the ground surface exploring
various crevices and burrows for suitable
spiders.

Twenty square-shaped transects were
established within the study site. The
dimension of each transect was 54 m2.
With the aid of several field assistants, I
walked through each transect in a linear
fashion, following paths delineated by
small yellow cords placed along the
ground in an east-west direction, and
separated by a distance of 6 m. Adult
wasps were collected using sweep nets.
We recorded the following data for each
wasp observed and collected: (1) time of
collection (Central Standard Time, CST);
(2) sex; (3) if feeding, the species of plant
(flower) being utilized; (4) type of plant for
wasps observed resting on vegetation; (5)
for flowers at which wasps were observed
feeding, flowers were collected and length
of the corolla tube was measured to the

Volume 15, Number 1, 2006

173

Table 1. Species of plants whose flowers were used as a source of nectar (percent utilization) by adults of
Pepsis grossa at a study site (Grassy Banks) located within Big Bend Ranch State Park, Brewster Co, Texas, during
2003. Data expressed as percentage utilization of 19 food resources. Data pooled for males (n = 504) and females
(n = 488). Scientific and common names of plants based on Powell (1988). B = Levin's measure of trophic
niche breadth.

Plant species

Mean corolla length (mm) (± SE)

Percent utilization

Ascepias texana Torrey (Texas Milkweed)

A. sperryi Woods (Sperry Milkweed)

Ungadia speciosa Endler (Mexican Buckeye)

Prosopis glandulosa Benson (Honey Mesquite)

P. pubescens Gray (Screwbean Mesquite)

Lycium pallidum Correll (Pale Wolfberry)

Diospyros texana Scheele (Texas Persimmon)

Agave lecluiguilla Torrey (Lechuguilla)

Yucca treculeaua Correll (Spanish Dagger)

Y. rostrata Engelman (Beaked Yucca)

Dasylirion texanum Scheele (Texas Sotol)

D. leiophyllum Engelman (Desert Candle)

Opuntia imbricata Hawes (Cane Cholla)

O. schotti Engelman (Dog Cholla)

O. phaeacantlia Engelman (Purple-fruited Pricklypear)

Salvia greggi Gray (Autumn Sage)

Forestiera angustifolia Torrey (Desert Olive)

Nolina erumpens Torrey (Bear grass)

Senecio douglasii Benson (Groundsel)

3.45 (0.41)
3.55 (0.23)
2.95 (0.23)
3.35 (0.28)
4.29 (0.18)
5.04 (0.31)
5.23 (2.45)
6.02 (2.06)

6.78 (3.03)
6.02 (1.87)
5.75 (1.93)
7.25 (2.11)
7.43 (2.04)
8.02 (2.18)

7.47 (1.99)

4.48 (0.32)
8.04 (1.97)
4.88 (0.67)

7.79 (1.08)

28.4
17.9
16.1
11.2

3.5

T.7

3.6

3.1

0.6

1.8

0.5

2.7

1.6

0.6

1.8

1.1

0.3

0.1
= 0.2816

nearest mm using a portable Unitron
dissecting microscope fitted with an ocular
measuring grid; (5) species of plants where
wasps were not observed to visit flowers.
Only those wasps that extended their
tongues into the corolla, or that were
observed to enter the corolla tube of
flowers with their entire head capsules
(and in some cases, part of the thorax as
well) and remain there for at least 20 sec
were considered to be in the act of feeding.
Each of these wasps was collected, frozen
on dry ice, and taken back to the laboratory
to confirm feeding by dissecting the gut
and analyzing gut contents.

Trophic niche breadth was determined
using the standarized Levin's index (B)
(Levins 1968): B = 1 /Y!,Pj> where pj re-
presents the proportion of individuals
(percent utilization) found on a particular
resource (plant species-flower). Values for
this measure can range from 0 (narrowest
trophic niche: all resources fall under one
resource category) to 1.0 (resources repre-
sented equally in all categories).

RESULTS AND DISCUSSION

During daylight hours, male and female
wasps were observed resting on shaded
areas of leaves or branches of Cottonwood
(Populus fremontii), willows (Salix taxifolia
and S. interior), walnut (Juglaus microcarpa),
oak (Quercus oblongi folia), mesquite (Proso-
pis pubescens and P. glandulosa), persimmon
(Diospyros texana), milkweed (Ascepias spe-
ciosa and A. speryii), Mexican buckeye
(Ugnadia speciosa), leatherstem (Jatropha
dioca), and soapberry (Sapiudus saponaria).
These same trees were used as perch sites
by males during April and May, when
activities associated with territorial defense
and breeding are most intense (Cazier and
Mortenson 1964, Punzo 2000, 2005).

The species of plants whose flowers
were used as a source of nectar by adults
of P. grossa are listed in Table 1. This
represents the first detailed list of specific
food plants for a wasp in this genus.
Because no significant differences were
found between males (n = 504) and

174

Journal of Hymenoptera Research

Table 2. Species of plants common at the Grassy Banks study site in Big Bend Ranch State Park whose
flowers were not observed to be visited and used as sources of nectar by adult males or females of Pepsis grossa
as sources of nectar. Scientific and common names of plants based on Powell (1988). Data on corolla tube length
(mm) expressed as means (± SE); N = number of each plant species.

Species

Aloysia gratissima Tronc. (Whitebrush)
Amsonia longiflora Torr. (Bluestar)
Cowania ericifolia Torr. (Heath Cliff Rose)
Croton doicus Cav. (Grassland Croton)
Euphorbia antisyphilitica Zucc. (Candelilla)
Fallugia paradoxa Endl. (Apache Plume)
Fouquieria splendens Engelm. (Ocotillo)
Hibiscus coulteri Harv. (Desert Rosemallow)
Justica warnockii Turner (Warnock Justica)
Krameria glandulosa Torr. (Range Krameria)
Lantana macropoda Torr. (Vinylleaf Lantane)
Mcndora longiflora Gray (Showy Mendora)
Leucophyllum frutescens Berl. (Purple Sage)
Selinocarpus parvifolias Standi. (Little Moonpod)
Senna wislizenii Gray (Senna)

Corolla tube length (mm)

617

10.05 (1.74)

448

9.02 (1.87)

194

7.79 (1.21)

847

5.81 (0.44)

524

5.06 (1.12)

456

8.05 (1.89)

905

13.84 (4.02)

389

7.52 (1.09)

197

5.94 (0.68)

295

4.35 (0.38)

428

13.21 (2.05)

236

14.56 (2.77)

683

11.93 (2.97)

379

16.84 (5.25)

257

4.76 (0.46)

females (n = 488) (Chi square test: X2 =
1.06, P > 0.06), data in Table 1 are
pooled for both sexes. Results indicate
that at this study site, adults of P. grossa
utilize the flowers of 4 out of 19 plant
species considerably more frequently
than the others, which is also reflected
in the value obtained for Levin's index.
It should also be pointed out that flowers
of the most frequently used species have
the shortest corolla lengths. These four
species accounted for 73.6% of the plants
whose flowers provided these wasps with
nectar.

In contrast, wasps were never observed
feeding from the flowers of other plant
species that were commonly found at this
site (Table 2). Ten of these 15 species have
flowers whose corolla lengths are in excess
of 7 mm, and 6 have values greater than
10 mm. In view of this, as well as from the
data in Table 1 on the four most-utilized
plant species, these results suggest that
adults of P. grossa prefer flowers with short
corolla lengths. However, a number of
species listed in Table 1 and 2, with corolla
lengths less than 6 mm, were visited in-
frequently or not at all by these wasps,
suggesting that other properties, in addi-

tion to corolla length, may influence
suitability of flowers as a food source.

According to a few previous reports,
Pepsis wasps from the southwestern United
States have been observed visiting the
flowers of a number of families of suffru-
tescent and woody flowering plants (Lin-
cecum 1867, Hurd 1948), although no
systematic attempt was made to identify
the species of possible food source plants.
Lincecum (1867) was the first to observe an
apparent preference of these wasps for
flowers of milkweed plants of the genus
Asccpias, which is in agreement with the
results of this study. In addition, these
wasps play an important role in the
pollination of milkweed plants (Hurd
1948).

In conclusion, the relatively narrow
trophic niche breadth exhibited by adults
of P. grossa, coupled with the fact that
flowers from all of the plants listed in
Tables 1 and 2 are common throughout the
spring and early summer at this study site,
indicates a preference for the flowers of
Asclepias species and Prosopis glandulosa. It
is interesting to note that flowers of
another mesquite species, P. pubesceus, are
visited far less frequently than those of P.

Volume 15, Number 1, 2006

175

glandulosa, even though these plants are
often found in close proximity. These
wasps may be using species-specific visual,
olfactory and /or gustatory cues to make
decisions as to which flowers to feed on.
Other species of nectivorous insects, in-
cluding various species of butterflies and
bees, are known to use combinations of
these types of cues to choose sources of
nectar (Heinrich 1979, Stone 1994).

ACKNOWLEDGMENTS

I thank J. Bottrell, K. Smart, L. Ludwig, P. Trepekan,
and B. Cummins for assistance in observing wasps
and recording data in the field, A. Simmons, G. Broad,
S. Cameron, and anonymous reviewers for comment-
ing on an earlier draft of the manuscript, and the
University of Tampa for providing me with financial
support (Faculty Development Grant) for this project.
Field studies were conducted with permission from
the Texas Dept. of Parks and Wildlife (Permit #: 41-
03).

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J. HYM. RES.
Vol. 15(1), 2006, pp. 177-180

NOTE

Polistes goeldii (Hymenoptera: Vespidae) is a Widespread but Rare

Social Wasp

Christopher K. Starr* and Allan W. Hook

(CKS) Dep't of Life Sciences, University of the West Indies, St Augustine, Trinidad & Tobago;

email: ckstarr99@hotmail.com
(AWH) Dep't of Biology, St Edward's University, Austin, Texas 78704, USA;

email: hook@acad.stedwards.edu

*Address for correspondence: Christopher K. Starr, Dep't of Life Sciences, University of the West

Indies, St Augustine, Trinidad & Tobago; tel (868) 662-2002 ext 3096 or 645-3232 ext 3096;

fax (868) 663-5241 or 663-9864; ckstarr99@hotmail.com

Key words: Polistes, neotropical, social wasp

Polistes goeldii Ducke was described from
the Amazon region of Brazil. It is a large,
robust social wasp, metallic blue-black
except for the mandibles and lower part
of the clypeus, which are reddish. Given its
size and color pattern, it bears a striking
resemblance to some members of the genus
Synoeca, especially the widespread S. sep-
tentrionalis Richards and S. surinama (L.).
Although P. goeldii can presumably deliver
a powerful sting, it has very small colonies
in comparison with the fearsome Synoeca
spp., so that the resemblance between the
two is almost certainly batesian mimicry.
That is, it seems out of the question that
Synoeca gains any defensive advantage
from potential predators' experience with
P. goeldii.

Polistes goeldii is a distinctive wasp with
an even more distinctive nest. In Septem-
ber 2004 we found an active colony of this
species at Caura Village, Trinidad, West
Indies with a nest matching in its overall
form all others that we have seen and that
have been described to us, as well as
a description by Richards (1978:522). It
was a single comb hanging from a highly
excentric petiole, attached to a fence wire
a little over one meter from the ground.
The petiole was relatively stout, about

3 mm long. The comb consisted of 22 cells
in just two rows, each cell attached some
distance below the base of its predecessor,
so that the comb descended sharply from
the petiole (Figs. 1-2). The largest nest of
this species of which we are aware had just
52 cells (Richards 1978:522).

The cell material was medium-gray
carton. The petiole and the comb top
around it were covered in shiny black
varnish, which was also applied in spots
elsewhere on the comb. Newer cells
showed less varnish than older ones,
consistent with the wasps periodically
applying it over the entire comb. The
pupal caps bulged moderately below the
cell mouths. They had only slight daubs of
pulp applied to them but were abundantly
spotted with dark varnish.

Over the course of three days we never
saw more than two adult females on the
nest, presumably the full number at that
stage. Three cells had fecal pellets in their
bases (Fig. 2), indicating that each had
produced an adult and now contained
a second brood individual. The walls of
these cells were cut back to make them
much shorter (Fig. 1).

Richards (1978) recorded P. goeldii from
a few localities in Brazil, Colombia, Ecua-

178

Journal of Hymenoptera Research

Fig. 1. Polistes goeldii colony in place at Caura
Village, Trinidad, West Indies. The length of the comb
is 128 mm. The upper wasp's mid- and hind-legs
straddle cells that have been cut back.

dor and Peru, as well as from "Salvador".
The identity of this latter is uncertain, but it
may refer to the city of that name in Bahia
state, Brazil. The published range of P.
goeldii, therefore, comprises an equatorial
belt over about 13 degrees of latitude on
both sides of the Andes.

In fact, the species is much more
widespread. Fig. 3 shows the presently
known localities of P. goeldii. These are
based on specimens that we have exam-
ined in the National Institute for Biodiver-
sity (INBio) in Costa Rica, Museum of the
Institute Agricultural Zoology (MIZA) of
the Central University of Venezuela, and
Land Arthropod Collection of the Univer-
sity of the West Indies (UWI) in Trinidad &
Tobago, and personal communications

Fig. 2. Cell map of the nest shown in Fig. 1. Cell
contents: 1 large larva (probably instar 4-5), o egg or
instar-1 larva, p with pupal cap, s small larva
(probably instar 2-3). The two lowest cells were
from J.M. Carpenter, J.H. Hunt, R.L. Jeanne empty. Asterisked cells have a fecal pellet. The petiole
and W.L. Overal, as well as Richards' was attached to the top-most cell.

Volume 15, Number 1, 2006

179

Fig. 3. Central America and northern South America, to show known localities of Polistes goeldii.

(1978) records with the exception of "Sal-
vador".

It is remarkable that the broad range of
such a distinctive wasp should have gone
unrecognized until now. The most likely
explanation is that P. goeldii is rare wher-
ever it occurs. As an example, at the time
we encountered the colony shown in Fig. 1
the authors had a combined residence of
about 16 years in Trinidad and had long
since come to suppose that we had
recorded all social wasp species existing
in the island. Yet this colony was nesting
just a few hundred meters from our
residence.

P. goeldii bears a strong overall resem-
blance to P. aterrimus Saussure, presum-
ably also a batesian mimic of Synoeca spp.

The two are most readily distinguished by
the pronotal keel (sharp and extensive in
each, but smoothly curving in P. goeldii,
versus with distinct humeral "shoulders"
in P. aterrimus) and the propodeal striae
(confined to the median furrow in P.
goeldii, versus extending strong onto the
sides in P. aterrimus), as well as by their
nests (P. aterrimus with a broad comb from
an excentric petiole, as in many other
neotropical species). The two species over-
lap very broadly in their ranges, although
there is a more or less distinct altitudinal
separation, P. aterrimus being found at
higher elevations.

Based on museum specimens, P. aterri-
mus appears to be much more common
than P. goeldii. In the INBio collection we

180

Journal of Hymenoptera Research

found just six specimens of P. goeldii,
versus 59 of P. aterrimus. The correspond-
ing figures for the MIZA collection are five
and 41. Similarly, the combined insect
museums of Colombia have four speci-
mens of P. goeldii and 32 of P. aterrimus
(C.E. Sarmiento, pers. comm.). In no case is
the preponderance of P. aterrimus speci-
mens due to a large nest series, so that the
comparison appears to be fair.

We have seen P. goeldii at Belem, Para,
Brazil, one of the type localities, and R.L.
Jeanne and W.L. Overal (pers. comms.)
report seeing it there as well. Even at
Belem, however, it appears to be uncom-
mon.

Across a broad range of plants and
animals, there is a clear positive correlation
between geographic range size and local
abundance (Gaston 2003:115-16). We are
not aware that this rule has been examined
with respect to any group of social insects,
but it is our definite impression that it
applies well to neotropical social wasps.
Such very widespread species as Polistes
versicolor (Olivier) and Polybia rejecta (F.),
for example, seem to be abundant almost
everywhere that they are found, while we
know of no very restricted species - with
the exception of some endemic to the
oceanic islands of the Antilles - that one
would characterize as common.

P. goeldii, then, appears to present
a striking exception to this rule. This
suggests that its niche is somehow un-
usually narrow for its genus. A good place
to look would be in its feeding habits.

The scant evidence available suggests
that colonies are characteristically small

and that nests never become large. It is
noteworthy that the Trinidad nest, com-
prising just 22 cells, had already produced
three adults and two pupae. A small
colony and nest proffer the possibility of
cryptic escape from predators, and several
features of the nest are consistent with such
an approach. The narrowness of the steep-
ly-hanging comb allows it to resemble
a short vine, something quite outside of
a predator's search image derived from
common species. This resemblance is en-
hanced by the application of dark varnish
on much of the comb. The removal of
carton from cells from which adults had
emerged may be an economy measure to
conserve material, or it may be a way of
diminishing the profile of the comb for
a time.

If predators and entomologists find P.
goeldi hard to locate, it is possible the
wasps themselves have difficulty finding
mates. It would be interesting to know if
this species has special adaptations in this
respect.

We thank the journal's reviewers (Sean
O'Donnell and Justin O. Schmidt) for
criticism of an earlier version of this note
and Azad Mohammed and Rajesh Ragoo
for help with the figures.

LITERATURE CITED

Gaston, K. J. 2003. The Structure and Dynamics of
Geographic Ranges. Oxford University Press, Ox-
ford.

Richards, O. W. 1978. The Social Wasps of the Americas,
Excluding the Vespinae. British Museum (Natural
History), London.

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